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State Biological Survey 


1^ University of Kansas 
% museum of natural history 




Miscellaneous Publication No. 5, pp. 1-68, plates 1-4, figures 1-15, 
^^(JJ & 1^ ^^^^^^^ March 10, 1953 






Introduction 4 

The Lesser Prairie Chicken 7 

The Greater Prairie Chicken 9 

History 9 

Characteristics of the Range and Present Distribution 12 

Life History of the Greater Prairie Chicken 17 

General Remarks 17 

Description of the Welda Area 18 

Daily Routine of One Flock 19 

Activities of Males on the Booming Ground 22 

Summer Activities of Males 23 

Reproductive Cycle of Females 23 

Nesting 24 

Rate of Egg Laying 25 

Period of Incubation 26 

Success of Nests 26 

Size of Clutch 27 

Decline in Size of Broods 29 

Plumages OF the Greater Prairie Chicken 31 

Development of Plumage in the Young 31 

Age Based on the Condition of the Molt 36 

Molt of the Adult Greater Prairie Chicken 38 

Length of Primaries as Correlated with Age and Sex 39 

Weights of the Greater Prahue Chicken in Autumn 41 

Food Habits of the Greater Prairie Chicken 43 

Population Changes of the Greater Prairie Chicken 51 

The 1949-1952 DecHne in Numbers 53 

^ Changes in Age Composition 56 

i Changes in Sex Composition 57 

) Differences in Abundance as Indicated by Hunters* Success . . 57 

^ Management 59 

'} Census 59 

Hunting Regulations and Refuges 61 

i Range and Pasture Management 62 

J Restocking 64 

'J Summary and Recommendations 64 

Literature Cited 66 




THE GREATER prairie chicken is a two-pound gallinaceous bird, 
of the grouse family ( Tetraonidae ) , formerly common but 
now rare or absent from many parts of its former range. Kansas, 
fortunately, still has extensive areas suitable for this bird, and is 
one of the four states having the largest number of the greater 
prairie chicken. The Dakotas and Nebraska are the other states. 
This bird is popular with hunters; other persons also take pride 
in the fact that sizeable flocks persist in parts of the State. But, 
what caused the bird to disappear in many areas when it remained 
in others? Can it be preserved? Can it be managed so that a 
part of the annual increase can be harvested as game, at intervals, 
without depleting the breeding stock? These and other ques- 
tions had occurred to many Kansans. At the University of Kan- 
sas, graduate students Lester Lew Henry and the late Wilbur S. 
Long had gathered considerable information concerning this game 
bird. Naturally, therefore, the greater prairie chicken was chosen 
for one of the initial studies when the State Biological Survey of 
Kansas was reactivated in 1949. 

Actually there are two species of prairie chickens and both occur, 
even today, in Kansas. The lesser prairie chicken, Tympanuchus 
pallidicinctus (Ridgway), lives in the southwestern prairies (see 
Figure 1). It is so uniform in size, color and bodily proportions 
throughout its range that no geographic populations are recognized 
as separate subspecies. 

The greater prairie chicken, Tympanuchus cupido (Linnaeus), 
occurs to the eastward of the other species (see Figure 1). Three 
geographic variants ( subspecies ) are recognized. The first of these 


is the heath hen, T. c. cupido (Linnaeus), that formerly occurred 
in the northern Atlantic States, but that now is extinct. The last 
bird of this subspecies died in 1931, and an account of the unsuc- 
cessful eflForts to save the subspecies is given by Gross (1928). 

110 100 90 80 70 

Fig. 1. Geographic distribution of the genus Tympanuchus Gloger. 
(1) Range of T. pallidicinctus (Ridgway); (2) original range of 
T. c. pinnatus (Brewster); (3) range occupied by T. c. pinnatus 
since cultivated crops were introduced; (4) former range of T. c. 
cupido (Linnaeus), now extinct; (5) range of T. c. attwateri 
Bendire. T. c. pinnatus now absent in much of its former range. 

The second subspecies is the Attwater prairie chicken, T. c. attwateri 
Bendire, which inhabits the prairies of the Gulf Coast of Texas. It 
still persists in a few places, and an account of its natural history 
is given by Lehmann (1941). The third subspecies is commonly 
known as the greater prairie chicken, T. c. pinnatus (Brewster); 
it occurs in the grasslands of the Midwest. 

Before the Midwest was settled the greater prairie chicken lived 
in the Tall-grass Prairies of the eastern and southern parts of what 


is now the United States. With the development of primitive agri- 
culture, it extended its range to the west and to the north as far as 
parts of the Prairie Provinces of Canada. As stated by Ridgway 
and Friedmann (1946:207), the range of the genus, originally and 
since the arrival of the white man, is "Open districts of eastern 
North America, from the western . . . Great Plains to the 
Atlantic coast ( locally ) and from Texas and southwestern Louisiana 
(formerly also Virginia?) northward to coast of Massachusetts, 
southwestern Ontario, southern Manitoba, and southwestern Sas- 

Ecologically, Kansas, with which we are immediately concerned, 
is an area of transition from the eastern hardwood forest to the 
western mixed prairie. Primitively, the diverse conditions in this 
transitional area provided suitable habitat for: the wild turkey, 
Meleagris gallopavo Linnaeus; the ruffed grouse, Bonasa umhellus 
(Linnaeus); the sharp-tailed grouse, Pedioecetes phasianellus (Lin- 
naeus); the bobwhite quail, Colinus virginianus (Linnaeus); the 
scaled quail, Callipepla squamata (Vigors); the lesser prairie 
chicken; and the greater prairie chicken. Without exception each 
of these birds, in Kansas, was at the periphery of its range. As 
Kansas was occupied and developed by white men from the eastern 
states, the turkey, the ruffed grouse and the sharp-tailed grouse 
became extinct in the State, and the ranges and numbers of the 
two prairie chickens were reduced. Recently residents of Rawlins 
and Cheyenne counties have reported that the sharp-tailed grouse 
is spreading from Nebraska into extreme northwestern Kansas 
(field notes on file in Univ. Kansas Museum of Nat. Hist., for 31 
October 1952, and 2 November 1952, R. H. Baker). Although 
man has introduced the ring-necked pheasant, Phasianus colchicus 
Linnaeus, to compensate partly for this loss, the fauna of gallin- 
aceous birds in Kansas is poorer by one species; this loss, at least 
in part, is the result of the activities of man. 

The aims of this study were to ascertain the present status of 
the prairie chickens in Kansas, and to learn previously imknown 
details of their life histories, habits and population behavior. Such 
information would be expected to aid in the conservation of the 
two species of birds. The investigation was concerned primarily 
with the greater prairie chicken, but such information as was ob- 
tained on the lesser prairie chicken is included. 

The co-operation and assistance rendered me by residents of the 
Welda Area where intensive field studies were made are gratefully 
acknowledged. Mr. William Brecheisen, Jr., was especially help- 


fill. Also I thank the employees of the Kansas Forestry, Fish and 
Game Commission, especially Mr. Dave Leahy, Director, for the 
assistance rendered in ascertaining the ranges of the two species 
of prairie chickens. These employees as well as several graduate 
students and faculty members at the University of Kansas gathered 
essential data on the numbers of the greater prairie chicken killed 
in the open seasons of 1950, 1951 and 1952. There at the University 
Mrs. Louise Brunk gave assistance with the line drawings. The 
drawings made by Mr. Richard Philip Grossenheider and Mr. Vic- 
tor Hogg are identifiable by their signatiu-es. Photographs are by 
the author, except as otherwise noted. Dr. Harrison B. TordoflF gave 
assistance in the study of the molt in the young of the greater 
prairie chicken, and Dr. Rollin H. Baker gave helpful criticisms, 
suggestions and encouragement throughout the course of the re- 
search. Both he and Dr. E. Raymond Hall gave critical assistance 
with the preparation of the manuscript. 

The Lesser Prairie Chicken 

The writers of many accounts concerning prairie chickens in 
Kansas do not differentiate between the two species. Therefore, 
and because the lesser prairie chicken was not recognized as a dis- 
tinct species until 1885, it is difficult to determine the early status 
of the lesser prairie chicken. Probably the chief breeding range 
of the lesser prairie chicken in Kansas always has been confined 
to the southwestern counties. According to Duck and Fletcher 
(1945?: 68), some early settlers in western Oklahoma recognized 
two kinds of prairie chickens in the same area, but occurring on 
different mating grounds. The *T)ooming and cooing kind" (greater 
prairie chicken ) was found in the uplands, and the "gobbling kind" 
(lesser prairie chicken) was found in the sandhills along water 
courses. If the former breeding range as mapped by Duck and 
Fletcher (op. cit.. Map II) in Oklahoma be extended into Kansas on 
the basis of similar soils, the former range would have extended as 
far east as the western part of Harper County, Kansas. 

In Kansas, the lesser prairie chicken has been taken as far east 
as Anderson and Neosho counties. The records from Neosho County 
are of a male and a female taken by Goss (see Goss, 1891:221) 
near Chanute on December 31, 1878, and on January 17, 1879, 
respectively. I have examined these mounted specimens and the 
original catalogue of Goss in the Kansas Historical Museum, To- 
peka, Kansas, and there is no doubt that these specimens are 
lesser prairie chickens. These records are incorrectly cited by 


Ridgway and Friedmann (1946:221) as from Neosho Falls, which 
was Goss' home. 

Other eastern records are as follows: A specimen in the Hurter 
Collection from southwestern Missouri, no date (Bent, 1932:285); 
a specimen in the collection of the Academy of Natural Sciences 
of Philadelphia from Garnett (spelled Garneth in Bent), Kansas, 
January 24-28, 1894 (Bent, loc. cit.); a male from Greenwood 
County, Kansas, in the "Rinker Collection," July or August 1895 
(Long, 1937:78). Probably it is significant that none of these 
specimens, for which the date of collection is given, was taken in 
the breeding season. The University of Kansas Museum of Natural 
History has only one specimen from outside the present breeding 
range of the species and that is from Logan County; the bird was 
taken on January 1, 1921. Game Protector E. L. Bryan reported 
to me {in litt., June 13, 1950) that the lesser prairie chicken formerly 
occurred in Trego, Ellis and Graham counties, Kansas. Probably 
this species formerly bred as far north as the counties just men- 
tioned, even though the greater prairie chicken also bred there. 

No evidence has been found that the lesser prairie chicken ever 
was reduced greatly in numbers in Kansas until the dry years of 
the 1930-1940 decade. Immediately prior to that time the species 
was abundant. According to the statement of a resident of Meade 
County, Kansas, the people of that area depended upon the lesser 
prairie chicken for food in place of domestic poultry, and the birds 
were taken whenever needed. In approximately 1928, three men . • 
shot 107 lesser prairie chickens on one morning, before 8 A. M., 
south of Garden City in Finney County. 

The drought of the 1930-1940 decade seems to have almost elim- 
inated the lesser prairie chicken in Kansas. Little food, cover or 
water was available over large areas, and numbers of lesser prairie 
chickens were reported to have been found dead with their nostrils 
clogged with dust. Edward Gebhard, of Meade, thinks that the 
only lesser prairie chickens left in Kansas at the end of the drought 
were on the XI Ranch, approximately 75 square miles, in Meade 
County, and a few on the Hitch Ranch in Seward County. "Buck** 
Adams of the XI Ranch reported that lesser prairie chickens came 
to the ranch headquarters for water in the drought years, and that 
only one small flock survived on the entire ranch. 

In 1950, the game protectors of the Kansas Forestry, Fish and 
Game Commission were questioned by mail to learn in which of the j 
105 counties prairie chickens were present. They were reported 
in 57 of the counties. Subsequent field investigation disclosed that 
the prairie chickens in the 14 southwestern counties were lesser 


prairie chickens, and that they were confined to the sandy lands 
that lie south of the Arkansas and Cimarron rivers (see Figure 2). 

The residents of southwestern Kansas report that these sandy 
lands supported stands of tall grasses before the drought of the 
1930-1940 decade. These grasses were ehminated over wide areas 
in the drought, and were replaced by sagebrush, Artemisia sp.; to 
date the grasses have not completely recovered. In ungrazed areas, 
tall grasses are crowding out the sagebrush, but in thousands of 
acres of rangeland, the sagebrush and short grasses predominate. 
Unless the native grasses effect a considerable recovery, the lesser 
prairie chicken may not approach its former abundance. 

Fig. 2. The geographic distribution of prairie chickens in Kansas. 
(1) Range of the lesser prairie chicken; (2) the chief range of 
the greater prairie chicken; (3) range wherein scattered flocks 
of the greater prairie chicken were reported present in 1950. 

A count of lesser prairie chickens on booming grounds on four 
and one-half square miles of some of the best range on the XI Ranch 
in the spring of 1951 revealed a total of 40 males on five booming 
grounds. A count on the same area in the spring of 1952 revealed 
82 males. This increase in numbers is encouraging, but the Hmited 
range suitable for the lesser prairie chicken in Kansas should be 
considered in any plans for its management. 

The Greater Prairie Chicken 

The distribution and abundance of this species in Kansas before 
the State was settled by white men, can only be inferred from the 


accounts of early explorers. It seems that the greater prairie chicken 
did not occur farther west than the middle of Kansas, and that the 
bird did not occur in impressively large numbers. The second 
point may be inferred from the lack of comment concerning the 
species by early explorers. Pike (see Coues, 1895:357-459) never 
mentioned seeing prairie chickens in his travels across Kansas in 
the autumn of 1806. Tixier (see McDermott, 1940:102-131) in 
1840 traveled overland from Independence, Missouri, to the Osage 
Village, thought to have been in Labette County, Kansas. He men- 
tions seeing prairie chickens (presumably greater prairie chickens) 
twice in Missouri near settlements, but never records their presence 
in the Territory of Kansas. More significantly, part of his party 
subsisted for two days on four upland plovers, Bartramia longicauda 
(Bechstein), during an enforced stay on the prairie. Prairie 
chickens probably were scarce or absent; if they had been abundant 
they probably would have been used for food in place of the upland 
plovers. Of approximately 25 references to the "prairie hen" in 
Thwaites (1904-1906), none is certainly referable to the genus 
Tympanuchus except in settled areas. Most significantly, Koch 
(1836:163) mentions that the numbers of prairie chickens (pre- 
sumably greater prairie chickens ) increased within three years after 
settlement of the prairie lands, and suggests that the cause of the 
increase was the food made available by cultivation. 

Baird (1860:628) wrote of the prairie chicken, "It scarcely seems 
to occur north of the United States line, nor, perhaps, beyond the 
beginning of the High Central Plains." McClanahan (1940:13) 
maps the western edge of the original range of this species in 
Kansas from approximately the eastern edge of Barber County, 
almost due north to the Nebraska-Kansas line. Duck and Fletcher 
(1945?: 68), after analyzing early records in Oklahoma and after 
talking to old residents of that state, concluded that the greater 
prairie chicken occupied most of the state east of the Panhandle, 
and mapped the western boundary of its range as crossing the 
Kansas line south of Medicine Lodge. Thus, it seems that before 
the coming of the white man, the greater prairie chicken in Kansas 
was confined to the eastern half of the State. 

Coincident with the development of intensive agriculture in the 
eastern part of the State, the numbers of greater prairie chickens 
declined in that area in the latter part of the 19th centmy. At 
the same time these birds occupied previously unused range in 
western Kansas, but later the center of population shifted back 
to the east. Goss (1891:225) says of this species, "common in the 


eastern and middle portion of the State and spreading westward 
with its settlement/' Cooke (1900:202) reported that the greater 
prairie chicken first nested in Colorado in approximately 1899. 
Dyche (1912:10) reports, "Informer years . . . prairie chickens 
were found in great numbers, especially in the eastern part. At 
present . . . prairie chickens are confined to counties in the 
western part of the State." Bunker (1913:146) states relative to 
the greater prairie chicken, "Formerly an abundant resident; still 
common in some parts of western Kansas." In the period 1912- 
1913, prairie chickens seemed to be at an all time low in Kansas; 
anyhow a few years later Clapp (1922:9) wrote: "Prairie chicken, 
one of the very finest game birds, and formerly abundant all over 
the State, was practically extinct ten years ago. This bird has come 
back handsomely and is now found in all sections where conditions 
are favorable, even in the extreme eastern counties of the State.** 

A parallel to this situation is described by Yeatter (1943:378) for 
Ilhnois. There, a general decline occurred from approximately 
1880 until 1903 when the hunting season was closed for the first 
time. By 1912, prairie chickens again had become sufficiently 
numerous in some areas to elicit complaints from farmers. Thus, the 
lowest population level in Illinois and Kansas did not occur at the 
same time, but both declines may have resulted from local condi- 
tions such as reduction in suitable range and overshooting. 

Since 1922, the greater prairie chicken has almost disappeared 
from northwestern Kansas, and has remained on a more or less 
stable range in parts of the eastern one-third of the State (see 
Figure 2). The record is not clear as to which species it was that 
Dyche (loc. cit.) and Clapp (loc. cit.) referred, but according to 
Long (1937:77) the remnant population in northwestern Kansas is 
of the greater prairie chicken. Specimens of the greater prairie 
chicken from western Kansas in the University of Kansas Museum 
of Natural History are, together with the year taken, from Ellis 
County, 1904; Rooks County, 1905; Trego County, 1906; Osborne 
County, 1907; Trego County, 1927. 

From the foregoing record, it seems that the population and range 
of the greater prairie chicken have changed much in the past 100 
years. Since the habitat requirements of a species would not be 
expected to change, the cause of such changes in occupied range 
must be sought in the habitat itself. Among possible causes of 
changes in habitat are changes in cHmate and land-use. Considering 
man as a part of the habitat, hunting pressure must also be con- 
sidered as a contributing factor to these changes. Flora (1948:3-5) 


says that there has been no definite trend toward increase or de- 
crease in amount of precipitation in Kansas, but that since ap- 
proximately 1890 there has been a trend toward warmer weather. 
This trend could hardly be a factor in the extension of the range 
of the greater prairie chicken into northwestern Kansas, because 
this bird can winter successfully in colder areas (Canada). 

In the biennial reports of the State Department of Agriculture, 
grain is mentioned as produced in Graham County as early as 1885 
and in Wallace County as early as 1890. The increase of winter 
foods, available to the greater prairie chicken, that accompanied 
this farming might well explain this westward extension of the 
range of the bird. It is thought that the changes in food and 
cover, especially the reduction of the tall grasses, that accompanied 
the dry years of the 1930-1940 decade, almost eliminated the greater 
prairie chicken from northwestern Kansas. 

The disappearance of the greater prairie chicken from much of 
eastern Kansas is attributable to the reduction of native grasslands 
by plowing and by the natural succession of woodlands after prairie 
fires were excluded. In areas where the greater prairie chicken 
now occurs in eastern Kansas, grasslands always have been at least 
as extensive as at present. The near elimination of the greater 
prairie chicken in eastern Kansas prior to 1913 may have resulted 
mostly from excessive hunting. 

Characteristics of the Range 
and Present Distribution 

Information received from game protectors in 1950 indicated that 
in four northwestern counties there were only scattered flocks of 
greater prairie chickens containing few birds. Greater prairie 
chickens in Washington, Clay, Ottawa, Saline, Ellsworth and Mc- 
Pherson counties also proved to be in widely scattered flocks using 
untilled parts of the Central Kansas Rolling Plains Region ( see Fly, 
1946: map). According to Fly (1946:163) much of this area is 
badly eroded. If needed soil conservation measures, including 
range — and pasture-improvement, were applied there, the area 
probably would become an important part of the range of the 
greater prairie chicken in Kansas. 

With the assistance of the local game protectors and other per- 
sons the principal range of the greater prairie chicken in Kansas 
was mapped as shown in Figure 3. Since Bennitt (1939:495) and 
Schwartz (1945:23) have ascertained that the greater prairie 
chicken is absent where there are no permanent grasslands, the 
presence of native grasslands was used to establish the margins of 


the range of this species in areas where the bird was reported to be 
present. The mapping all was initially done in the field on county 
maps with a scale of one-half inch to one mile. 

Fig. 3. The principal range of the greater prairie chicken in 
Kansas. (1) The Western Area; (2) the Bluestem Hills; (3) the 
Eastern Area; (4) the Blackjack Prairie. Scattered flocks occur 
in other counties included in the range in Figure 2. 


The principal range of the greater prairie chicken is classified 
into four types, each a modification of Fly's (1946) Natural Agri- 
cultural Resource Areas of Kansas ( see Figure 3 ) . Data concerning 
farm crops from townships representative of each type for 1950 
were obtained from the files of the Kansas Crop and Livestock 
Reporting Service, 203 Federal Building, Topeka, Kansas. These 
data were gathered by township assessors and were from civil 
townships varying in size from 13,000 to 66,000 acres. Table 1 
lists the minimum, average and maximum per cent of the total 
farmland in native grass and in feed crops for each range type and 
for six townships in southeastern Kansas which are considered to 
be marginal range for the greater prairie chicken. The term "feed 
crops'' as used here includes corn, wheat, sorghum and soy-beans. 
These four crops provide the chief foods used by the greater prairie 
chicken in autumn and winter. 

Table 1. The Per Cent of the Total Farmland in Native Grass and in 

Feed Crops 


Feed Crops 


Average Maximum 











Prairie . . 

. . 58.17 





















. 37.69 






The names of the subdivisions of the principal range of the 
greater prairie chicken in Kansas, as hereinafter used, were chosen 
by me as descriptive of the location or chief characteristic of each 
subdivision. The **backbone" of the bluestem prairie country in 
Kansas is a narrow strip extending from the Oklahoma border in 
Chautauqua and Cowley counties to near Marysville in Marshall 
County. This area is divisible into two parts. The southeastern 
part, the Blackjack Prairie Area, is characterized by the presence 
of woods on some hilltops. The remainder of the area has almost 
no woody cover on the uplands and is designated as the Bluestem 
Hills Area. To the east and west of these areas the percentage 
of the total farmland that is in native grass becomes progressively 
less with increasing distance. Some areas, extending to the eastern 
and western borders of the principal range of the greater prairie 


chicken in Kansas, are designated as the Eastern Area and the West- 
em Area respectively. See figure 3 on page 13. 

The Eastern Area (Plate 1, Figure a). — This type includes parts 
of Fly's (op. cit.) East Central Prairies, the eastern part of his 
Bluestem Hills and the eastern part of his Cross Timbers and 
Interspersed Prairies. The Eastern Area is typified by a mixture 
of croplands and native grasslands; the grasslands are in excess 
of fifty per cent of the farmland. Croplands are rarely more than 
one mile from any part of the grasslands; consequently there is 
a favorable interspersion of winter food and cover. The burning 
of pastures is practiced but, because roads and croplands act as 
barriers, fire usually is limited to small areas. This type of range 
includes the best range for the greater prairie chicken in Kansas. 

Blackjack Prairie (Plate 1, Figure h). — This is the central part 
of the area designated by Fly (op. cit.) as "Cross Timbers and 
Interspersed Prairies." The soils are sandy, and there are woods on 
many hilltops. Areas of prairie are extensive and in some places are 
farther removed from feed crops than in the Eastern Area. 

Bluestem Hills (Plate 1, Figure c). — This type includes only 
the central part of Fly's (op. cit.) area of the same name. Soils 
are flinty or cherty, cultivation is limited to the creek valleys, and 
areas of grassland may be several miles across, resulting, in many 
places, in poor interspersion of food and cover. Burning of pastures 
in the spring is common, especially on leased land, and large con- 
tinuous areas are burned. In the spring of 1950 I drove from Casso- 
day to Cottonwood Falls, a distance of 24 miles, in this range type 
without seeing any unbumed grassland. The greater prairie chicken 
was seen twice on this same trip. Spring burning and poor inter- 
spersion of winter food and cover are primary limiting factors for 
the greater prairie chicken in this large area. The Bluestem Hills 
differ from the Blackjack Prairie in having more extensive grass- 
lands, no woody vegetation on the upland, and in having a cal- 
careous parent material for the soil. Unless further investigation 
reveals that these two areas differ markedly in the abundance of the 
greater prairie chicken, they might be considered as a unit when a 
management plan is put into effect. 

Western Area (Plate 4, Figure a). — This type is the transition 
between the Bluestem Hills and the intensively cultivated land 
to the west. The agricultural characteristics resemble those of the 
Eastern Area, except that the grass-flora of the uplands, when 
grazed, tends strongly toward the short- and the mid-grasses. 

Marginal Townships. — Six townships, where the greater prairie 


chicken occurs in small numbers, were studied to ascertain the 
per cent of feed crops and of grasslands in areas of marginal quality 
for this bird. These findings appear in Table 1. 

Optimum Townships. — The areas of greatest abundance of the 
greater prairie chicken in Kansas are in southwestern Anderson 
County, southwestern CoflFey County and northwestern Woodson 
County. Two townships characteristic of the best of this range 
were studied to determine the pattern of land use that is optimum 
for the greater prairie chicken. The summary of land use is as 

Anderson County, Welda Township 

Per cent in grass 62.87 

Per cent in feed crops 18.64 

Woodson County, Center Township 

Per cent in grass 66.35 

Per cent in feed crops 16.64 

It is evident that approximately one-third of the land must be 
in permanent grass to provide the minimum requirements for the 
greater prairie chicken in Kansas, and that approximately two- 
thirds in permanent grass provides the optimum condition. The 
findings relative to the minimum requirements are in close agree- 
ment with those of Schwartz (1945:23) for Missouri, but he found 
no correlation above the minimum between the amount of grassland 
and the number of prairie chickens. It will be shown later that the 
Bluestem Hills Area supports fewer prairie chickens than does the 
Eastern Area. This fact is thought to be the result, at least in part, 
of the poor interspersion of food and cover in the Bluestem Hills 
Area in Kansas. See figure 3 on page 13. 


Life History of the Greater Prairie Chicken 

General Remarks 

Earlier workers, notably Bent (1932) and Schwartz (1944 and 
1945), have discussed the life history of the greater prairie chicken 
in detail. In the autumn, flocks or packs assemble and function as 
a unit in their daily movements throughout the ensuing winter. 
These packs may be all of one sex or of both sexes. With the ad- 
vent of warmer weather — anytime from late January to early 
March — the males separate themselves from the females, and be- 
gin to visit places, known as booming-grounds, where territorial dis- 
putes and courtship displays take place each morning and evening. 
The duration and intensity of these disputes and displays depend 
to some extent on the condition of the weather. In late March 
the females begin to visit the booming grounds, and usually in the 
first half of April a peak of mating activity is reached. 

Nests are made in dry vegetation left from the previous season. 
Some sets of eggs hatch before mid-May, but most of them hatch 
in late May or early June, after an incubation period of 22 or 23 
days. Late nests, which are re-nestings after failure of an eariler 
attempt, hatch as late as July 18, but there are few of these as com- 
pared with those that hatch in May and June. 

The males continue courtship activities until hot weather, usually 
until early June. Pursuit of the females is not hmited to the 
booming grounds, but occurs wherever the females are found. 
As attendance at the booming grounds wanes, the flocks of males 
disintegrate. The resulting singles and small flocks spend their 
time loafing about swales and other places where suitable shade 
is provided by shrubs and tall grass. Females that are not success- 
ful in bringing off a brood follow the same routine. 

The females that succeed in bringing off a brood stay with their 
young all summer, and frequent both permanent grasslands and 


croplands. By the first of September, the young are indistinguish- 
able from the adults, except when the birds are in the hand, and at 
approximately this time they begin to assemble in flocks. 

In the autumn of 1949 a part of Welda Township, Anderson 
County, was chosen for the intensive study of the life history of 
the greater prairie chicken. Originally it was planned to use ap- 
proximately one township for this study, but the density of the 
population of prairie chickens, and other factors, made it more 
practical to select a small area where a single flock could be studied 
in greater detail. The arrangement of roads, fields and farmsteads 
and the positions of ponds, booming grounds and feeding areas in 
the Welda Area are shown in figure 4. 

Description of the Welda Area 

Permanent grasslands used both for pasture and for hay are of 
native species, predominantly the bluestems, Andropogon scoparius 
Michx. and A. gerardi Vitman, the latter occurring mostly in un- 

FiG. 4. The Welda Area. Twp. 22S, R. 19E, Anderson County, 
Kansas. C — cropland; G — grassland used for hay; Gp — grassland 
used for pasture; F — ^farmyard; F-1 and F-2 — feeding areas; B-1, 
B-2, etc. — booming grounds. Ponds are represented by irregular 
enclosed areas and the center of sections by circles. 

grazed areas. In most pastures, grazing is well regulated, and good 
stands of grass occur, but in the northeast quarter of section 15, 
and about the large pond in section 10, overgrazing by cattle is 
severe and annual weeds, predominantly ragweeds. Ambrosia ar- 
temisiifolia L. and A. bidentata Michx., blue grass, Poa pratensis 


L., and annual grasses such as foxtail, Setaria sp., form approxi- 
mately one-half of the ground cover. 

In addition to the areas marked as meadow in Figure 4, much 
of the grass in the well-drained parts of the west half of section 10 
was cut for hay each summer. The grassland of sections 9 and 10 
was burned each spring in March. No other burning occurred. 

The only trees in section 11 and in the north half of section 14, 
other than an abandoned farm grove and a plum thicket in the 
northwest quarter of section 11, are six mulberry, Morus rubra L., 
three small white elm, Ulmus americana L., and one cottonwood, 
Populus deltoides Marsh. A hedgerow of Osage orange, Madura 
pomifera Raf., and other hardwoods and cedar, Juniperus virginiana 
L., extends along the west side of the road between sections 10 and 
11, and along the north side of the field designated as F-2. There 
is another hedge of Osage orange along the west side of the pasture 
in the northwest quarter of section 15. Other woody growth in the 
western part of the area consists of widely scattered mulberry trees, 
cottonwoods and elms, and shrubs along the waterways. These 
shrubs are predominantly false indigo, Amorpha fruticosa L., and 
buttonbush, Cephalanthus occidentalis L. 

Sources of water were plentiful, because rainfall was normal or 
above normal each summer, and pools of water occurred at fre- 
quent intervals in all waterways. 

Cultivated crops in 1949, 1950 and 1951 were sorghums of dif- 
ferent varieties, corn, wheat, oats, soybeans and alfalfa. White 
clover and Korean lespedeza occurred in the pastures. Each cul- 
tivated area was subdivided, and crops were rotated within each 
such area. As a result the same feed crops were available in each 
area used by the prairie chickens for feeding each year. The north- 
west quarter of section 15 changed ownership in 1949, and has been 
more intensively farmed since that year. Terraces were built in 
the large cultivated area of this quarter in the fall of 1950. 

Approximately 22 per cent of the area of study was in cultivation, 
77 per cent in grassland and the remainder of the area in farmyards. 

Daily Routine of One Flock 

The flock (hereinafter referred to as Flock A), resident in the 
western part of the Welda Area, was observed periodically through- 
out the period of study, November 8, 1949 to May 1, 1952. Obser- 
vations were made from a blind and from an automobile at boom- 
ing grounds and feeding places with the aid of 7 X 50 binoculars 
or a 20 power telescope. Records were kept of all observed flock- 
movements. Color banding of birds trapped and released, and 


subsequent sight records and returns supplemented these obser- 
vations. Further information was obtained concerning roosting 
places and general activity of the birds by walking through the 
area. Flock A was composed entirely of males and contained 145 
birds in the fall of 1949; it declined to 15 birds by the spring of 1952. 

In the fall and winter of 1949-1950, the daily routine of Flock A 
began with a flight from the roosting grounds on unmown slopes 
in the southwest quarter of section 10. This flight to booming 
ground number one ( B-1 ) usually occurred approximately one-half 
hour before sunrise, but was later on days with inclement weather. 
After a few minutes to an hour there, where some birds would feed 
in the adjacent cornfield, the flock would fly west to the cultivated 
land in the northeast quarter of section 16 (F-1), where the prin- 
cipal morning feeding would be done on waste cane and soybeans. 

Fig. 5. The usual ranges of four flocks of greater prairie chickens 
using the Welda Area in winter, and the movements of banded 
birds (cf. Fig. 8). Banding sites are indicated by dots and the 
movements of banded birds by arrows. 

After the flock fed, the birds would return to the vicinity of the 
roosting area, or to a similar site, where the middle of the day would 
be spent loafing. In the evening, the feeding area and the boom- 
ing ground would again be visited, but not in so regular an order 
as in the morning. The birds returned to the roost usually when it 
was too dark to observe them well, except against the western sky. 

The routine of this flock in 1950-1951 was of the same pattern, 
but booming ground number two (B-2), and the feeding area in 
the southeast quarter of section ten (F-2) were used to the exclu- 


sion of those formerly used. These changes in daily habits of this 
flock concurred with the terracing of B-1 and the 1950 hunting 
season. Each year the range of daily activity of Flock A covered 
approximately one square mile, but the composite range for the 
two yeais was approximately one and one-half square miles. The 
flocks using the Welda Area are shown in figure 5. 

To the southwest, outside th^ study area, there was another 
booming ground. Birds from the vicinity of this booming ground 
(Flock B) fed, in part, at F-1 with Flock A, but the two flocks 
were distinct at other times. Birds from the vicinity of B-4 and 
B-5 (Flock C) fed with Flock A at F-2 but the chief range of Flock 
C was to the south where it fed outside the area. Another flock 
( Flock D ) came from the east to feed at F-2. This flock was thought 
to be composed of females; as many as 20 females were seen at 
F-2 at one time. This was the approximate size of Flock D. 

Trapping proved to be extremely difficult because of the mild 
winters and abundant food; the greater prairie chickens seldom 
came to the bait placed to entice them into traps. Even so, ten birds 
were trapped and banded. One of these was taken in a tip-top trap 
(see Hamerstrom, 1942:7), four were taken in a projected net 
trap (see Dill and Thornsberry, 1950), and five in hoop net traps 
powered by rat traps of the snap-type. Colored bands and num- 
bered aluminum bands were used in combination so that each 
banded bird could be identified if observed at close range. Sight 
records on booming grounds or returns from dead birds, or both, 
were obtained from eight of these ten birds. The movements of 
individual birds, made evident by these records, are shown in 
Figure 5. Bird number one, a male, was trapped at F-1 on De- 
cember 28, 1949, and was seen at B-1 on March 24, 1950. Birds 
number two and three were banded at B-1 on October 14 and 19, 

1950, respectively. One of these two was seen at B-3 on April 25, 

1951. Number four was banded at F-2 on December 16, 1950, 
and was seen at B-1 on April 24, 1951. Number five was banded 
at the large pond in the south part of section 10, on December 28, 

1950, and was seen repeatedly at B-2 in the spring of 1951. Num- 
bers six and seven were banded at B-2 on March 27 and 30, 1951. 
Neither returned to B-2, but both were seen at B-3 on April 25, 

1951. Number seven was shot approximately 200 yards east of the 
banding site on October 25, 1951. Numbers eight, nine and ten 
were banded at F-2 on April 7, 1951. Number nine, the only fe- 
male banded, was found dead within 200 yards of the banding site 
on May 3, 1951. Number eight was shot less than one-half mile 


east of the banding site on October 24, 1951. Number ten was 
not seen after banding. No banded birds were observed at boom- 
ing grounds in the spring of 1952. 

These observations agree with the observations of flock be- 
havior and together they indicate that different flocks share the 
same feeding ground, but act as distinct units at other times, and 
that the daily and seasonal ranges of flocks and individuals were 
limited to a cruising radius of approximately one-half mile. 

These observations stand in contrast to those in Wisconsin of 
Hamerstrom and Hamerstrom (1949) who found winter packs to 
be made up of several smaller flocks. These packs operated as 
units in severe weather but subdivided into smaller groups in mild 
weather. The ranges of the various packs covered 2-4 square 
miles and did not overlap. Schwartz (1945:83) found similar be- 
havior in the greater prairie chicken in Missouri. There, flocks of 
both sexes banded into packs and cruised over most of the range 
of the separate flocks. In Nebraska, Mohler (1952:22) found the 
home range of flocks in winter to be 2000 acres ( approximately three 
square miles) or more. These differences in behavior of flocks 
in wdnter may be attributed to the mild winter weather that pre- 
vailed in the course of this study, and to the greater density of popu- 
lation in the Welda Area. 

Activities of Males on the Booming Ground 

The activities of the greater prairie chicken on booming grounds 
have been so completely described by Schwartz (1944 and 1945), 
and others, that only the observed seasonal changes in activity will 
be presented here. The earliest autumnal activity on a booming 
ground was observed three and one-half miles southwest of Ottawa, 
Franklin County, on September 15, 1949, where 24 males assembled 
in the morning, engaged in territorial disputes, and unsuccessfully 
attempted to boom. At the time of the first visit to the Welda Area, 
on November 8, 1949, approximately 100 males were at B-1 and 
territorial disputes and booming were in full progress. Attendance 
at booming grounds was noted at the time of each morning visit 
by the observer that autumn and vdnter (1949-1950), and in the 
next autumn and winter (1950-1951). Schwartz (1945:58) found 
that the greater prairie chicken in Missouri did not visit booming 
grounds in mid-winter. 

Hamerstrom (1939:108) concluded that activity on booming 
grounds was determined by upper and lower limits of a combina- 
tion of light and temperature. This conclusion explains the mid- 
winter booming-ground activity observed by me in 1949-1950 and 


1950-1951, since both winters were mild compared to those when 
Schwartz made his study. The operation of the upper threshold of 
light and temperature is indicated by the latest observed booming- 
ground activity on June 12, 1951, which was a cloudy, cool day. As 
late as May 30, 1951, spirited booming-ground activities and indica- 
tions that hens were present on the booming ground were noted. 

Summer Activities of Males 

In summer, during the molt, males seldom are seen, and they 
are reluctant to fly when disturbed. Adults of both sexes were ob- 
served in low areas where shrubs and associated tall grasses pro- 
vided favored loafing cover. No evidence was found that the few 
upland trees were used as loafing cover by adults in summer. 

Reproductive Cycle of Females 

In winter, too few females were observed to justify drawing 
definite conclusions as to their activities then. The greater prairie 
chicken was seen in many places other than the Welda Area, and 
in all instances males predominated or no females were seen. It 
seems that the daily routine of females involves fewer conspicuous 
movements by flight than does that of males. 

Although no intensive study of the behavior of females in the 
breeding season was attempted, such observations as were made 
merit the same conclusions as those drawn by Schwartz (1945:51). 
In his discussion of the spring booming-season he concludes, "Dur- 
ing the early part of the season the principal activities on the 
booming ground are territorial disputes and booming. No females 
are seen there until late in March, when an occasional hen visits 
the booming ground for a short time. . . . These infrequent 
visits may continue for a week or more; then suddenly the num- 
ber of hens visiting on a booming ground increases to a rather 
constant maximum, marking for several days or a week the so-called 
height of the season. . . . After this period the number of hens 
decreases very rapidly to a few who come in only occasionally." 

In this study, the earliest observation of a female on a booming 
ground was on February 25, 1951. The peak of mating, in 1950, 
occurred in the second week in April. In 1951, although seven 
mornings were spent observing booming-ground activity from a 
blind between March 24 and April 25, no peak of mating activity 
was observed. The largest number of females observed in any one 
morning, in 1951, was three on April 13. 

Hamerstrom (1939:112) concluded that most of the nests of the 
greater prairie chicken in Wisconsin were begun at the height of 


the booming-season. Yeatter (1943:385), on the other hand, found 
that breeding in IlHnois was spread more uniformly over a period 
of time and concluded that this was because all of the birds did 
not attain sexual readiness at the same time. 

Schwartz (1945:53) observed that males remain in their in- 
dividual territories, on the booming ground, and allow matings in 
adjacent territories to proceed without interruption only at the 
height of the breeding season. At other times, males frequently 
interrupt matings in adjacent territories by driving the male from 
the back of the female. This is not to imply that all matings occur 
at the height of the season, for courtship and matings occur at 
places other than the booming ground, and the spread of hatching 
dates, including those of re-nestings, indicates that matings occur 
over at least a six-week period of time. Both sexes seem to be- 
come physiologically incapable of breeding shortly after the first 
of June. This characteristic is most significant, because it limits the 
reproductive period to late spring and early summer, and makes 
the success of reproduction largely dependent upon the favorable 
weather within this short period. Bobwhite quail, on the other 
hand, remain paired throughout the summer, and have been known 
to hatch young as late as October. Frequently the main hatch 
of bobwhites occurs in July. In Missouri, the peak of hatching in 
quail was in the first half of July in 1948, in the last half of July 
in 1949, and between July 7 and August 7 in 1950 (see Stanford, 
1950 and 1951 ) . The value of the ability thus to persist in nesting is 
obvious, in that it tends to insure a high reproductive rate even 
though weather conditions may be unfavorable for nesting and rear- 
ing young in the first part of the summer. 


To find nests and broods of the greater prairie chicken, a flushing 
device designed after that of Lehmann (1946) was made to be 
used on a light truck, tractor or jeep. The essential feature of this 
device was a drag suspended from a horizontal bar that was 
mounted on the vehicle. The horizontal bar was made of steel 
pipe in three sections, each ten feet long, the outer two of which 
telescoped into the center section. A central post provided means 
of supporting the ends by wires. It was found necessary to use 
springs in the supporting wares to prevent breakage. The most 
satisfactory drag was made of pieces of steel pipe the same length 
as the sections of the horizontal bar. Figure b of Plate 4 illustrates 
this device mounted on a truck. Later, two sections were added 
making a total spread of 46 feet. This bar on a hght tractor was 


satisfactory. In one instance 160 acres were censused in six hours. 

From May 12 to June 19, 1951, 16 nests were found in 610 acres 
of unburned pastures and meadows. No broods indicative of earlier 
nests were found, and no nests were found later than June 19 while 
I was searching for broods with the drag. This tract of 610 acres 
was given complete coverage and it is thought that all nests were 
found, because the number of successful nests and the number of 
broods found agreed closely. 

The value of studying nests is sometimes questioned, because of 
the effects of the disturbance caused by the study. For example, 
three nests were destroyed by my efforts to find them. My study, 
otherwise, however, had little effect on the success of nests. No 
nests were approached on foot, but only in a vehicle. This prac- 
tice should have minimized any tendency on the part of predators 
to follow human trails. Hens were difficult to flush from the nest, 
and in one instance the hen did not flush until the wheel of the 
truck passed over her tail and pulled out all rectrices and coverts. 
This hen returned to the nest and succeeded in bringing off her 
brood. In another instance a hen permitted the wheel of the truck 
to pass within one foot of one side of the nest and the inner end of 
the drag to pass within one foot of tlie other side without flushing. 
In a third instance, the nest was destroyed by a predator only after 
the hen had returned and laid two more egss. 

Rate of Egg Laying 

Little direct evidence is available as to the rate at which prairie 
chickens lay eggs. Gross (see Bent, 1932:248) found in nests of 
captive greater prairie chickens and in one nest of a wild bird that 
the period of laying was approximately twice as long in days as 
the number of eggs laid. Lehmann (1941:15) reported that the 
same species (T. c. attwateri) in Texas normally laid one egg 
per day until the clutch was complete, but that sometimes there 
were intervals of one to three days between the times of egg-laying. 
Indirect evidence obtained in this study proves that in some in- 
stances egg-laying is at the rate of one egg per day. Two nests 
found on May 29, 1951, each contained seven unstained eggs 
plus some stained eggs. The stained eggs had been laid, or were 
present, in a rainy period. The unstained eggs had been laid 
after the rains ceased. Examination of the weather data for the 
lola weather station, for which hourly readings of precipitation were 
available, revealed that rain occurred each day from May 16 to 23 
inclusive and ended early in the morning of May 23. A light 


shower recorded at lola on the 26th did not extend to the study 
area. If each of these hens had laid one egg per day from May 23 
to 29 inclusive, each would have laid seven eggs. 

Period of Incubation 

Schwartz (1945:66) observed two nests of the greater prairie 
chicken from the beginning until the end of incubation. The eggs 
were incubated 23 and 24 days. In my study a nest that was found 
on May 24, 1951, contained eggs that were incubated at the time 
of discovery and that hatched 22 or 23 days later. 

Success of Nests 

Assuming that each hen laid one egg per day, and that the period 
of incubation is 23 days, the date on which the first egg was laid 
was calculated for each nest. One or two eggs from each nest 
were tested in water to determine the stage of incubation (see 
Westerskov, 1950 ) . For nests that did not hatch, the date on which 
the first egg was laid was estimated by this means. The fate of 
each nest is described below. The nests are arranged in the order 
of the estimated date on which the first egg was laid. All dates refer 
to 1951. 

No. 1. Discovered on May 25, 13 eggs, hit with tractor wheel; salvaged 
four eggs, one hatched in incubator on May 29. 

No. 2. Discovered on May 25, 15 eggs; on June 3 all had hatched at 
latest by the day before. 

No. 3. Discovered on May 25, 13 eggs, hit with tractor wheel; salvaged 8 
eggs, hatched in incubator June 1. 

No. 4. Discovered on May 25, 13 eggs, hit with tractor wheel; salvaged all 
eggs, hatched in incubator June 1. 

There was one infertile egg among the 39 from nests 1, 3 and 4. 

No. 5. Discovered on May 25, 15 eggs; hatched June 6. At 8 A. M. on 
June 6 the hen was on the nest with the complete brood. The disturbance 
caused by my marking several of the chicks, the subsequent trampling about by 
calves, and a heavy rain in the forenoon caused a loss of chicks as foUows: 
one chick trampled in the nest; two stepped on by calves; one dead but no 
obvious physical damage; one caught by the foot in heavy grass, alive but 
barely able to move; one sitting quietly near the nest. The two that were 
ahve were taken and successfully reared. The chilled one recovered within 
20 minutes after being placed inside my shirt. These two were picked up at 
3:15 P. M. when a hen was heard but not seen near the nest. 

No. 6. Discovered on May 28, 13 eggs; all hatched on June 3. 

No. 7. Discovered on May 26, one egg was cracked; the remaining 13 all 
hatched on June 9; one hve chick left behind in the nest. 

No. 8. Discovered on May 24, 12 eggs; all hatched June 16. 

No. 9. Discovered on May 24, 8 eggs; all eggs gone without evidence of 
cause on June 6. 

No. 10. Discovered on May 25, 7 eggs; female seen on the nest May 26; 


nest had been destroyed on June 3 when the remains of 9 eggs were found. 
The cause of this loss is uncertain, but tooth marks left on one shell corre- 
sponded in width to that between the canine teeth of the raccoon, Procyon 
lotor Linnaeus, or a small dog. According to Stoddard (1931:188), dogs 
usually eat the entire egg, whereas these eggs had but one side removed in a 
manner characteristic of the raccoon (see Reardon, 1951). 

No. 11. Discovered on May 29, 11 eggs; found destroyed on June 3, canine 
marks and a tuft of hair on one egg indicated that a spotted skunk, Spilogale 
interrupta Rafinesque, had destroyed the nest (see Plate 4, Figure c). 

No. 12. Discovered on May 29, 9 eggs; all gone on June 3 without evi- 
dence of cause. 

No. 13. Discovered on May 28, 2 eggs; found destroyed on June 3, ends of 
eggs removed and fragments carried oflF 15 feet from the nest in a manner 
characteristic of the cotton rat, Sigmodon hispidus Say and Ord, as described 
in Stoddard (1931:140). 

No. 14. Discovered on June 16, 9 eggs. On the night of June 29-30 there 
was a rain of 2.12 inches at lola (2.33 inches at Gamett). At 11 A. M. on 
June 30 I visited this nest and found no evidence of the hen; all but two eggs 
were pipped, at least part of the chicks were alive in the shell. At 4 P. M. 
the hen was at the nest, and one egg was hatching. On July 2, the shells 
of two hatched eggs were found here; the other seven had failed to hatch. 
This nest was situated on the steep side of a natural drainageway. The hen 
probably was driven from the nest at hatching time by water. 

No. 15. Discovered on June 19, 9 eggs; on June 30 all were hatched; the 
shells were stained and the egg and embryonic membranes were separated from 
the shells which suggested that they had hatched several days prior to June 30. 

No. 16. Discovered on June 14 while I was searching the same area covered 
on May 25, 9 eggs; never located again; thought to have been destroyed. 

Hamerstrom (1939:114) reported that in Wisconsin 50 per cent 
of the nests were successful. In Illinois, Yeatter (1943:392) found 
49 per cent of the nests to be successful. Seven of 16 nests found 
in this study were successful. Five of these seven were among the 
first eight when arranged according to the date when the first egg 
was laid. The remaining three of the first eight are those de- 
stroyed by the tractor, all of which were in an advanced stage of 
incubation, and at least part of which might be presumed to have 
been successful if undisturbed by me. While it is true that all 
other unsuccessful nests had incomplete clutches or were in early 
stages of incubation when found, none of these nests was known 
to have been abandoned before destruction. The successful nests 
were found on the first or second, eighth, thirteenth, fourteenth and 
seventeenth days of incubation, as judged by the dates of hatching. 

Size of Clutch 

The size of full clutches reported by other observers ranges from 
a minimum of five (see Hamerstrom, 1939:111) to a maximum of 
25 (see Schwartz, 1945:65). The largest clutches were thought 


by Bent (1932:248) to be the product of two or more hens. The 
maximum clutch thought to have been produced by one hen is 17 
(see Hamerstrom, loc. cit.). The average size of 66 clutches in 
Wisconsin was 12 (Hamerstrom 1939:113). 

Audubon (1834:494) noted that second nests had fewer eggs 
than first nests. Hamerstrom {loc. cit.) found that there was a 
decline in the average size of the clutch as the date on which the 
first egg was laid was progressively later. A similar dechne in size 
of clutch was noted in the 13 full clutches found in 1951 on the 
Welda Area. Figure 6 shows the temporal distribution of these 
clutches, their size and the regression line of size of clutch against 
date of first egg laid. The regression coefficient, r = — .702, rep- 
resents a high degree of negative correlation between the date of 
first egg laid and the size of the clutch. 











25 30 5 10 15 20 25 30 4 



Fig. 6. Scatter diagram of size of clutch (Y axis) plotted against 
estimated date on which first egg was laid (X axis) and the regres- 
sion hne of Y on X. r = — .702. Data from the Welda Area, 1951. 

The dates on which first eggs were laid occur in three groups 
(see Figure 6). The mean intervals between these groups are 
16 and 17 days. This grouping suggests that re-nesting occurred 
after failure of first nests. Hamerstrom (1939:115) found that the 
greater prairie chicken has certain breeding characteristics in com- 
mon with the bobwhite quail, which is known to re-nest, and con- 
cluded from this that the prairie chicken re-nests. Lehmann (1941: 
15) found that some Attwater prairie chickens re-nest as many 
as two times, after failures, for a total of three nesting efforts. 

In summary, although the sample is small, it seems clear that 
early clutches are larger, are more successful, and produce most 
of the young. Sixty-eight of the 79 chicks that were produced from 
16 nests came from the earUest eight nests. As Lehmann ( 1941:15) 
says relative to the Attwater prairie chicken, "A successful season 


depends largely on the fate of the early nests, so that a primary 
objective of management should be to safeguard these attempts."* 

Decline in Size of Broods 

In 1950, when I was unable to be on the study area, Mr. William 
Brecheisen, Jr. recorded all broods seen in the course of his normal 
farming operations. He did not estimate ages of the chicks after 
they were half grown, and the data that he gathered can best be 
treated according to the date of observation. The results of his ob- 
servations, presented in Figure 7, reveal that broods ranged in size 
from three to fifteen, averaged approximately eight chicks at the 
beginning of the season and averaged 6.6 at the end of the season 
in August. 



ec 10 

















Fig. 7. Scatter diagram of size of brood seen (Y axis) plotted 
against the date of observation (X axis) and the regression line of 
Y on X. r = —.191. Data from the Welda Area, 1950. 

In 1951, the flushing device previously described was used to 
find broods in pastures; and early morning work with a dog located 
broods in cultivated land where the flushing device could not be 
used. Six hundred and twenty acres of the study area, plus eighty 
acres outside the area, were censused. A total of eighteen broods 
able to fly were found ranging in number of individuals per brood 
from two to twelve. Figure 8 presents the distribution and decline 
in size of these broods. The average number of individuals per 
brood was four and one-half as compared with approximately seven 
in 1950. The decline in number of individuals per brood, after the 
young were able to fly, was less than in 1950, r = — .091 in 1951 as 
compared to r = — .191 in 1950. Another difference between the two 
seasons was that the first broods were seen 19 days earlier in 1950. 

If it is true that breeding activity is controlled by a combination 
of light and temperature (see Hamerstrom, 1939:108), then a com- 


parison of the average temperature and cloudiness, as reflected 
by the total precipitation, in the two winters might help to explain 
the 19-day delay in hatching in 1951. Presumably, the three 
months before the normal onset of breeding activity would afiFect 
this phenomenon. In 1950, the temperature was 1.3 degrees above 
normal for this period and the precipitation was 2.02 inches below 
normal (normal is 4.37). In 1951, the temperature was 2.7 degrees 
below normal and the precipitation was 0.82 inches above normal for 
the same period. Thus, the combination of light and temperature 
was presumably less in 1951. While a study over a much longer 
period of time, employing more refined techniques for measuring 
the amount of Hght, would be necessary to justify drawing any 
definite conclusions, these data do indicate that the winter weather 
in 1951 might have delayed the onset of breeding. 



• • • • • 

I ' ' I I I I I I I I'll 

10 20 30 40 50 60 70 


Fig. 8. Scatter diagram of size of brood seen (Y axis) plotted 
against estimated age of brood in days (X axis) and regression line 
of Y on X. r = —.091. Data are from the Welda Area, 1951. 

The smaller size of the broods observed in 1951 probably was a 
result of the unusually heavy precipitation. Schwartz (1945:67) 
called attention to the destructive eflFects of heavy rain in the hatch- 
ing period. In 1951, the average size of clutches in successful nests 
was 12 eggs; nevertheless at the time when the chicks were able 
to fly, approximately ten days of age, the average brood contained 
less than five chicks (approximately eight in 1950). Precipitation 
in June of 1951 was 12.04 inches at Gamett. This is 7.30 inches 
above normal for this part of eastern Kansas. Precipitation and 
temperature in the same period of 1950 were near normal. 

In summary, in most years, and possibly in every year, the most 
critical time for the greater prairie chicken in Kansas seems to be 


when nesting and rearing of the young occur, namely from the 
middle of May until the middle of July. 

Plumages of the Greater Prairie Chicken 
Development of Plumage in the Young 

Little is known of the details of the molt in prairie chickens, 
because only in rare instances have prairie chickens been suc- 
cessfully reared in captivity whereby birds of known age were avail- 
able for study. Lehmann ( 1941 : 16 ) gives an account of the growth 
of the Attwater prairie chicken based on observations of wild birds. 

Dwight (1900:164) observed that the postjuvenal molt in prairie 
chickens is complete except for the two outermost primaries. The 
condition of these two primaries was used by Ammann (1944) in 
diflFerentiating between birds of the year and adults in autumn 
populations. Petrides and Nestler (1943 and 1952) developed 
techniques for aging young bobwhites by noting the condition of 
the molt of the remiges. This technique is a valuable tool in de- 
termining the time of the major hatch in wild birds — an important 
determination in the study of population dynamics. 

Twenty-five eggs of the greater prairie chicken, salvaged from 
nests 1, 3 and 4, were placed in an incubator at 103° F. Seventeen 
chicks hatched from these eggs and were placed in an electrically 
heated brooder. The newly hatched chicks fed readily on insects, 
including mealworms, and some learned to eat commercial chick 
feed. Others did not learn to eat, and seven died by the fourth day 
after hatching. Moistened chick mash and lettuce leaves were the 
staple foods of those that survived after the fourth day. 

The legs and joints of some birds that survived after the fourth 
day became swollen (Plate 2, Figure d). In some individuals the 
use of one leg was lost. After the fourth day, birds were sacrificed 
at intervals to the 29th day. In the meantime, two chicks taken 
from nest number 5 were placed in the brooder, and later were 
reared. One died at the age of 18 weeks and the other one was 
released at the point of capture at the age of 20 weeks. 

Insofar as possible these captive birds were examined once each 
week, but after the twelfth week it was not possible to handle them 
without undue risk of injury to them. In each examination, the 
length of each remex was taken by placing the end of a rule against 
the insertion of the feather in the skin and flattening the feather 
against the rule. Supplementary information was obtained from 
seven wild-taken juvenal greater prairie chickens. The condition 


of the molt was noted for each and the remiges were measured ac- 
cording to the same system as was used for the captive birds. 

Pterylosis of the greater prairie chicken is essentially the same 
as that of the ruffed grouse as described by Trainer and Holm 
(see Bump et al., 1947:76-90 and 741). Terminology used herein 
is the same as that used by Trainer and Holm except that the pri- 
maries are numbered from proximal to distal, the terms supra-ocular 
and infra-ocular are used to denote the areas immediately above 
and below the eye and that the term "cervical apteria*' is used 
to designate the bare areas on the sides of the neck. 

Because the number of birds available for study was small it is 
impossible to infer the probable amount of variation that occurs 
between individuals, and thus to determine the extent to which the 
data may be used in estimating the ages of birds in the field. 
However, the results do form a tentative scale for aging immature 
greater prairie chickens. In the following descriptions, emphasis 
is placed on the changes that occur as the birds become older. All 
measurements of primary feathers and secondary feathers are given 
in numerical order. The number of each specimen is the catalogue 
number of the University of Kansas Museum of Natural History. 
The unnumbered specimens are the two captives. 

Fig. 9. Condition of the remiges of the greater prairie chicken at 
hatching. Sheaths of juvenal primary feathers 1-7 and their 
greater upper coverts are present. Sheaths of juvenal secondary 
feathers 3-13 and juvenal greater upper coverts 1-13 are present. 
All other remiges are present as natal down, and down feathers 
remain attached to the tips of the sheaths. 

Twelve hours. No. 30473 KU (Figure 9). Natal down throughout, ex- 
cept on the alar tract where there are, of the juvenal plumage, primaries 1-7, 
greater primary coverts 2-7, secondaries 3-13 and greater secondary coverts 
1-13; all juvenal feathers are sheathed and are of approximately same length, 
7 nmi. 


Thirty-six hours. No. 30474 KU. Sheaths of primaries extend beyond the 

Sixty hours. No. 30475 KU. Primaries 2-7 unsheathed at tips; greater 
primary coverts unsheathed at tips; secondaries and their coverts still sheathed; 
juvenal feathers of alula present and sheathed. 

Sixty-six hours. No. 30492 KU. No appreciable change from condition 
at sixty hours. 

Eighty hours. No. 30476 KU. Lengths of first seven primaries 6, 9, 10, 
9, 8, 6, 4 mm.; juvenal scapulars present and sheathed. 

Eighty-five hours. No. 30477 KU. Primary one unsheathed at tip; lengths 
of primaries 9, 10, 10, 11, 12, 11, 7 mm.; greater coverts cover base of pri- 
maries; lesser primary coverts present and beginning to unsheath; juvenal 
feathers of alula unsheathed (visible in Plate 2, Figure b). 

Four days. No. 30479 KU. Lengths of primaries 9, 10, 10, 11, 11, 12, 10 
mm.; secondaries unsheathed at tips. 

Six days. No. 30478 KU. Lengths of primaries 20, 22, 28, 28, 28, 26, 22 
mm.; juvenal scapulars unsheathed at tips. 

Eight days. No. 30482 KU. Lengths of primaries 25, 30, 30, 32, 30, 27, 
23 mm.; first juvenal feathers of sternal region present, sheathed; neither 
juvenal scapulars nor sternal feathers yet visible through undisturbed down. 

Ten days. No. 34083 KU. Lengths of primaries 33, 36, 37, 38, 39, 33, 29, 
11 mm.; primary eight present for first time; juvenal scapulars unsheathed at 
tips; most advanced juvenal feathers of sternal region imsheathed at tips; 
superior margins of sternal region and of femoral tract with quills of juvenal 
feathers (visible in Plate 2, Figure c). 

Eleven days. No. 30484 KU. Lengths of primaries 38, 39, 40, 38, 40, 
38, 33, 9 mm.; juvenal feathers of dorsal cervical region and of interscapular 
region now present and sheathed: appearance of new juvenal feathers in 
sternal region progressing ventrad. 

Thirteen days. No. 30485 KU. Lengths of primaries 50, 51, 52, 52, 49, 
48, 40, 20 mm.; no appreciable change otherwise. 

Fifteen days. No. 30486 KU. Growth complete in juvenal primaries 1-3, 
lengths as at 13 days, lengths of others 57, 54, 50, 46, 20 mm.; a few sheaths 
of juvenal feathers on coronal region; in cervical region, sheaths of juvenal 
feathers present from base of neck to posterior border of cervical apteria; 
juvenal feathers of interscapular region unsheathed and concealing down; 
sheaths of juvenal feathers of posterior dorsal region present, but hidden in 
down; juvenal rectrices present and unsheathed, juvenal feathers of sternal 
region in paired patches, the sheathed feathers meeting at midline (see Plate 2, 
Figures c and d). 

Twenty-two days. No. 30487 KU. Lengths of primaries 48, 62, 60, 66, 
71, 71, 65, 42 mm.; primaries 9-10 visible in flesh; a few unsheathed juvenal 
feathers on midline of coronal region; juvenal auriculars present; juvenal 
dorsal cervicals extend to above cervical apteria; juvenal feathers of posterior 
dorsal region cover down; juvenal rectrices average 28 mm. long; juvenal 
undertail-coverts equal rectrices; juvenal ventral cervicals ex-tend to below 
cervical apteria; sternal region covered with juvenal feathers except at midline 
(see Plate 2, Figure c). 

Twenty-nine days. No. 30488 KU. Juvenal primary 1 replaced; primary 
one sheathed, 14 mm. long; lengths of primaries 2-10 are 60, 72, 82, 86, 90, 
82, 68, 25, 13 mm.; coronal and occipital regions with juvenal feathers of 


characteristic russet color; auriculars pronounced; juvenal cervicals present 
to anterior border of cervical apteria; juvenal restrices average 45 mm. long; 
juvenal plumage complete on sternal region and femoral tract; crural and 
pedal tracts with sheathed juvenal feathers throughout (see Plate 2, Figure /). 

The following descriptions are based on notes made from captive birds and 
deal primarily with postjuvenal molt of wing. 

Forty-one days. Male. Lengths of first-winter primaries 1-2, 77 and 32 
mm.; first-winter primary number 3 visible in flesh; juvenal primaries 4-8 
completely grown; primaries 9-10, 64 and 36 mm. long; juvenal pinnae 
slightly longer than other cervicals. Female. Primaries in same stage of 
replacement as in male; first-winter primaries 1-2, 71 and 45 mm. long; 
juvenal primaries 9-10, 52 and 39 mm. long; both specimens with plumage of 
head slightly advanced over that of 29-day-old birds, but natal down still 
evident; no postjuvenal molt on body (see Plate 3, Figure b). 

Forty-eight days. Male. First-winter primaries 1-4, 87, 73, 62, 20 mm. 
long; juvenal primaries 5-10, 89, 96, 109, 115, 86, 62 mm. long; first-winter 
secondary number 3, 29 mm. long; other secondaries juvenal. Female. Molt 
of primaries same as in male. Lengths of primaries 82, 93, 57, 17, 113, 115, 
124, 128, 86, 65 mm.; molt of secondaries same as in male; secondary number 3, 
32 mm. long ( see Plate 3, Figure c ) . 

Fifty-five days. Female. First-winter primaries 1-5, 97, 97, 78, 60, 
l5 mm. long and all growing; juvenal primaries 6-10, 92, 108, 122, 102, 70 mm. 
long; 9-10 growing; first-winter secondaries 3-4, 52 and 37 mm. long; all other 
secondaries juvenal and grown (see Plate 3, Figure d). 

Sixty days. Male. First-winter primaries 1-5, 96, 108, 108, 88, 52 mm. 
long; number 6 missing; juvenal primaries 7-10, 115, 132, 113, 87 mm. long; 
first-winter secondaries 3-5, 86, 64, 20 mm. long; greater secondary coverts 
1-8 of first-winter plumage. Female. Molt of primaries same as in male. 
Primaries 93, 97, 92, 77, 45, 0, 100, 120, 100, 83 mm. long; primary coverts 
1-5 replaced; first-winter secondaries 3-5, 75, 55, 15 mm. long; greater 
secondary coverts 1-7 of first-winter plumage. 

Sixty-seven days. Female. First- winter primaries 1-6, 98, 102, 110, 112, 
87, 44 mm. long; juvenal primaries 7-10, 100, 121, 128, 98 mm. long; 9-10 
growing; first-winter secondaries 3-6, 82, 48, 38, 10 mm. long and growing. 

Seventy-five days. Male. First-winter primaries 1-7, 96, 102, 120, 136, 
110, 96, 20 mm. long; juvenal primaries 8-10, 110, 146, 116 mm. long; 9 and 
10 growing. 

Eighty-two days. Male. First-winter primaries 1-8, 105, 124, 128, 140, 
165, 160, 110, 30 mm. long; juvenal primaries 9-10, 152 and 125 mm. long; 
secondary 1 juvenal; 2-10 first-winter and nearly grown. Female. First- 
winter primaries 1-7, 100, 105, 111, 121, 143, 128, 67 mm. long; 8 missing; 
juvenal primaries 9-10, 137 and 115 mm. long and fully grown; secondary 1 
juvenal; number 2 first-winter and 38 mm. long; 3-4 first-winter and grown; 
5-8 first-winter and 86, 78, 56, 26 mm. long and growing; other secondaries 
juvenal (see Plate 3, Figure e). 

One hundred and twenty-six days. No. 30472 KU. Male. Primaries 105, 
114, 125, 142, 170, 178, 176, 166, 152, 124 mm. long; 1-8 first-winter, 9-10 
juvenal; only number 8 growing; secondaries all replaced and 88, 102, 101, 
101, 99, 99, 96, 96, 94, 94, 92, 91, 88 mm. long; all other tracts in complete 
first-winter plumage but with growing feathers throughout. 



Fig. b. Oak thicket in extensive grassland in the Blackjack Prairie. Eight miles 
\ Fall Riv., Greenwood Co., Kan., July 14, 1952. Photo No. 45 KIMNH. 

Fig. c. l:.\U'nM\i' p.i.sUut' land txpual ot tin liliicsle'iu llills. i)i\v iiiili' west 
of Eskridge, \\'al)aunsee County, Kansas, Jul\ 13, 1952. Photo Xo. 46 KUMXH. 


Fig. b. Flushing-bar mounted on a State Biological Survey truck. Welda 
Area, June 13, 1951. Photo No. 48 KUMNH. 

Fie. c. Nest No. 11 after destruction b\ a spotted skunk. "Welda Area, 
June 3, 1952. Photo No. 51 KUMNH. 

The sequence of molt of the body plumage can be described more accurately 
from freshly killed specimens than from Hve birds, or from dried skins. The 
condition of the molt was noted in seven freshly killed young greater prairie 
chickens in the summer of 1951. The notes made from these birds form the 
basis for the following descriptions. 

No. 30490 KU. Male. Molt of primaries slightly advanced over that of 
birds forty-one days old. First-winter primaries 1-3, 95, 66, 21 mm. long; 
juvenal primaries 4-10, 89, 96, 99, 110, 106, 54, 40 mm. long; primaries 1, 
2, 3, 8, 9, 10 growing; secondaries all juvenal; interramal, infra-ocular and 
supra-ocular regions with natal down; pedal tract with sheathed juvenal 
feathers; all other regions juvenal. 

No. 30469 KU. Female. Primary replacement same as in female fifty-five 
days old. First-winter primaries 1-5, 106, 112, 97, 62, 11 mm. long; juvenal 
primaries 6-10, 111, 122, 133, 112, 87 mm. long; first-winter secondaries 3-4, 
31 and 5 mm. long; other secondaries juvenal; capital tract juvenal throughout; 
cervical and interscapular regions with first-winter feathers interspersed with 
juvenal feathers; no replacement in posterior dorsal region; three central rec- 
trices missing ( thought to be accidental ) ; ventral tract with first -winter feathers 
throughout, not showing beyond juvenal feathers except at superior margin of 
sternal region; femoral and crural tracts with first- winter feathers as sheaths, 
not visible beyond juvenal feathers. 

No. 30470 KU. Male. A sick bird from same brood as No. 30469 KU. 
First-winter primaries 1-5, 105, 105, 109, 93, 36 mm. long; juvenal primaries 
6-10, 115, 125, 128, 109, 80 mm. long; first-winter secondaries 3-4, 79 and 
36 mm. long; otherwise as in No. 30469 KU except all rectrices present. 

No. 30493 KU. Male. Primary replacement same as in two birds sixty days 
old. First-winter primaries 1-5, 107, 115, 112, 84, 41 mm. long; primary 6 
missing; juvenal primaries 7-10, 126, 140, 125, 89 mm. long; primaries 3, 4, 
5, 9, and 10 growing; first-winter secondaries 3-5, 83, 34, 10 mm. long; capital 
tract juvenal; first-winter sheaths present on cervical region and scapular tract; 
posterior dorsal region with first-winter feathers except posteriorly; no replace- 
ment in caudal tract; first-winter feathers visible on ventral tract from between 
cervical apteria to anterodorsal sternal region and to femoral tract; some first- 
winter sheaths hidden in juvenal feathers on midline anteriorly but not 

No. 30495 KU. Male. Primary replacement same as in male seventy-five 
days old. First-winter primaries 1-7, 112, 118, 118, 130, 111, 66, 10 mm. long; 
juvenal primaries 8-10, 127, 136, 114 mm. long; primaries 5, 6, 7, 9 and 10 
growing; secondary 1 juvenal and 83 mm. long; secondary 2 missing; first- 
winter secondaries 3-9, 113, 116, 83, 65, 53, 36, 21 mm. long; secondaries 5-9 
growing; greater primary coverts 1-7 replaced; greater secondary coverts all 
replaced, grading in condition of growth from fully grown distally to un- 
sheathed at tip only proximally; under primary and secondary coverts present 
as sheaths; capital tract and anterior cervical region all juvenal, otherwise 
spinal tract with first-winter feathers throughout; two central rectrices missing 
(thought to be accidental); first-winter stemals meet at midline anteriorly 
but not posteriorly; abdominals, crurals and pedals juvenal. 

No. 30489 KU. Male. Primary replacement same as in female eighty-two 
days old. First-winter primaries 1-7, 114, 116, 120, 140, 164, 150, 90 mm. long; 
primary 8 missing; juvenal primaries 9-10, 161 and 156 mm. long; primaries 
6-7 growing; secondary 1 missing; first-winter secondaries 2-13, 117, 110, 


110, 110, 108, 106, 102, 89, 77, 65, 45, 24 mm. long; capital tract with first- 
winter feathers on coronal region, juvenal feathers elsewhere; dorsal and ventral 
cervical feathers of first-winter plumage to anterior border of cervical apteria, 
remainder of spinal tract completely of first winter feathers except juvenal 
feathers interspersed in interscapular region; rectrices juvenal; upper tail- 
coverts of first-winter feathers; undertail-coverts of juvenal feathers; ventral 
tract with first-winter feathers meeting at midhne anteriorly, extending pos- 
teriorly along superior margin to femoral tract; abdominal region with a few 
first-winter feathers posteriorly, otherwise with juvenal feathers (see Plate 3, 
Figure /); femoral and crural tracts predominantly of first-winter feathers; 
pedals juvenal. 

Summary of Molt. — ^The postnatal and postjuvenal molts occur 
in the same sequence. Juvenal primaries 1-7 are present at hatch- 
ing and subsequently 8, 9 and 10 appear in that order. Juvenal 
primary number 1 is molted at four weeks of age, and at tbat age 
juvenal primaries 9 and 10 appear. Molt of the primaries proceeds 
distally, and on the average one feather is molted in each wing 
each week. Number 8 is molted at twelve weeks of age. Numbers 
9 and 10 are retained through the first winter. The greater primary 
coverts are replaced in the same sequence and approximately at 
the same time as the corresponding primaries. 

Secondaries 3-13 are present at hatching, and all are present 
in the juvenal plumage at the age of six weeks. The postjuvenal 
molt of the secondaries begins with number 3 and proceeds proxi- 
mally, but with more variation between individuals than there 
is in the order of appearance of the primaries. Secondaries 2 and 1 
are lost in that order when the postjuvenal molt of the primaries is 
nearing completion at approximately ten weeks of age. 

At the time of hatching, the entire body, head and legs, excepting 
the apteria, are covered with natal down. As a rule, in a molt, the 
first feathers to appear in a given region are in the anterior or dorsal 
part of that region, and growth proceeds posteriad and ventrad. 
An exception to this is the cervical plumage which appears first 
on the posterior part of the neck. Posterior dorsal feathers and 
abdominal feathers appear over the entire region at one time. The 
first feathers to appear on the body in a molt are the scapulars; 
they are followed by the stemals, interscapulars and the femorals. 
The capital, caudal and pedal tracts are the last to molt. The 
rectrices seemingly are molted all at one time, or nearly so, since no 
progressive loss of tail feathers is evident in any specimens at hand. 

Age Based on the Condition of the Molt 

Molt in the young of the greater prairie chicken is an orderly 
process. On four occasions, greater prairie chickens, too young 




Fig. 10. Condition of the molt of the primaries of the greater 
prairie chicken one week to twelve weeks of age. Blackened bases 
of the feathers indicate that the feather is growing; juvenal feath- 
ers are stippled and first-winter feathers are cross-barred. Feather 
number one is at the right in each series of ten primaries. 


to fly far, were captured and examined in the field. Comparison 
of these individuals and the specimens collected in the wild with 
the captive birds, demonstrates that captives correspond closely 
with wild birds both in growth and in development of their plum- 
age. Comparison of the wild birds with the captive birds permits 
the investigator to age the wild birds accurately enough to deter- 
mine the time of the peak of hatching. 

As an aid to ascertaining the age of young prairie chickens, charts 
illustrating the development of the primaries are presented in 
Figure 10. Only data from captive birds of known age were used 
in preparing these charts. For each primaiy an outline of the 
fully formed adult feather is shown. Within this outhne, the con- 
dition of each feather is shown at the age under consideration. A 
feather which is growing, is indicated by the base of the quill being 
blackened. Juvenal feathers are shown as stippled, and first-winter 
feathers are shown as cross barred. The photographs (Pis. 2 and 
3) illustrate the condition of the molt of the body at various ages. 
These photographs are of captive birds, except Plate 3, figure /. 
These figures demonstrate the close correspondence in body plum- 
age of wild and captive birds with identical condition of molt of 
the wing. 

Molt of the Adult Greater Prairie Chicken 

Adult males begin to molt after the cessation of breeding activity. 
At this time they become secretive and reluctant to fly. No females 
with broods were collected, but those without broods that were 
examined corresponded to the males in the progress of molt. 

No. 30494 KU. Male. Taken August 25, 1951. Molt of wing 
and body feathers complete; rectrices not molted; capital and pedal 
tracts not molted; pinnae replaced and 35 mm. long. 

No. 29914 KU. Male. Taken September 15, 1950. All primaries 
molted, 9-10 present as sheaths; all secondaries molted, number 1, 
26 mm. shorter than number 2; rectrices not molted; sheaths of 
new feathers on throat. 

No. 30491 KU. Female. Taken July 25, 1951. Primaries 1-6 
replaced, number 6 a sheath; corresponding greater coverts re- 
placed; secondaries 3-4 replaced; number 4 a sheath; new feathers 
on coronal region; interscapular region with approximately 30 per 
cent new feathers; posterior dorsal region not replaced; ventral 
tract with new feathers ahead of wing, otherwise not replaced; 
caudal tract not replaced. 

No. 20 M. F. B. Female. Taken on July 28, 1951. Primaries 
1-5 replaced; secondaries not replaced; capital tract and anterior 


cervical region with only sheaths of new feathers; caudal tract as 
follows: two outer left, four outer right and central left rectrices 
old; all others new and of greater length medially. This is the only 
specimen examined with partial molt of the rectrices. 

Summary. — The general pattern of the postnuptial molt seems 
to be the same as that of the postjuvenal molt, especially of the 
remiges. The body molt starts near the base of the wings, but 
may occur rapidly and may be general over the body, with the 
capital, caudal and pedal tracts being last as in the young. 

Length of Primaries as Correlated with Age and Sex 

Measurements were taken of primary number eight of each bird, 
killed by hunters, examined in the open season, on October 24, 
1951. This was to ascertain the amount of variation in the length 
of this feather and to determine whether or not its length might 
be used as a criterion of sex. Since the age class (adult or im- 
mature) can be ascertained by the condition of the two outer 
primaries, both sex and age could be ascertained from the wing 
alone if it were possible to separate the sexes by this means. 
Measurements were taken in milhmeters by placing the end of a 
rule against the hind edge of the wing at the base of the feather and 
flattening the feather against the rule. Data were obtained from 206 
birds, and were recorded with the classification of each bird as to 
sex and age. To aid in the interpretation of these data, wings 
were saved from 23 birds, and measurements were made of both 
the seventh and eighth primaries of these wings. 

Table 2. Length Characteristics of Primary Feathers of the Greater 

— s 

Prairie Chicken (cf. Fig. 11). X±2 —is the 0.9546 Confidence Limits 


of the Mean; Sx is the Standard Deviation; V is the Coefficient of Varia- 
tion; n IS THE Number of Individuals in the Sample. Extrexces of Length 
ARE in Millimeters. 

Extremes _ « 

of X ± 2 — Sx V n 

Length " 

Primary 8 141-172 159.30 ± 3.74 8.99 5.64 23 

Primary 7 153-171 162.00 ± 2.54 5.45 3.77 23 

All males 138-178 166.29 ± 1.40 7.08 4.26 102 

Selected males 151-178 168.89 ± 1.14 5.72 3.43 99 

All females 98-178 158.41 ± 1.26 6.42 4.05 104 

Selected females 148-178 159.18 ± 1.10 5.51 3.41 100 

AU adults 142-178 163.78 ± 1.34 7.11 4.34 112 

All young 98-178 162.55 it 1.68 8.13 5.00 94 


Variation in the length of the eighth primary was found to occur, 
and this primary had not completed its growth in some of the birds 
examined. To compare the variation due to incomplete growth 
with the variation due to individual differences, primaries seven 
and eight, from the 23 wings, were compared. 

Table 2 and Figure 11 summarize the analyzed data from all the 
wings that were examined, both in the field and in the laboratory. 
In the table the data headed "primary 7" and "primary 8" are from 
the 23 wings examined in the laboratory, and the other data are 
presented by classes of age and sex. The presentation in Figure 11 
is by "Dice squares" which include the range, mean, one standard 
deviation either side of the mean and two standard errors either 
side of the mean. Two standard errors either side of the mean 
represent the 0.9546 confidence limits that may be placed on a 
given sample. Thus, where these do not overlap, the difference 
between the means is significant at that level. The coefficient of 
variation is a measure of the variability — the smaller the coeffi- 
cient the less the variabihty. 

In the 23 wings, the seventh primary is less variable than is the 
eighth, V = 3.77 as compared to 5.64. This difference is caused 
by the shortness of the eighth primary in some birds; this short- 
ness is indicated by the longer range below the mean in Figure 19. 
The pulpy bases of the shafts indicate that these feathers are not 
completely grown. Contrasted with the data from primary number 
eight, those for number seven are completely symmetrical (see 
Figure 11) indicating a normal distribution of primary lengths. 

In samples from the field, the lengths of some feathers stood 
apart from those of others on the lower end of the scale, indicating 
incomplete growth in those feathers. It seemed reasonable that 
the data might be refined by dropping these measurements from 
consideration when comparing lengths of primaries of males and 
females. This was done, and these categories are indicated as 
selected males and selected females. It is apparent (see Figure 
11) that the average length of primary eight is different between 
males and females, and that this difference is statistically significant 
at a high level of confidence, but that the range in length is too 
great to permit the separation of sexes by this means, even when 
the lower extremes (incompletely grown feathers) are eliminated 
as in the instance of the selected groups. The average length of 
primary number eight is not significantly different between birds- 
of-the-year and adults in the samples available to me and it is 
thought that a larger sample would reveal no significant difference. 















■■ I — 



■' I ■ " 


140 150 





Fig. 11. Variation in the length of primary feathers of the greater 
prairie chicken by sex and age (cf. Table 2). The horizontal line 
indicates the range, the vertical line indicates the mean, the open 
rectangle indicates one standard deviation either side of the mean 
and the blackened rectangle indicates two standard errors either 
side of the mean. 

Weights of the Greater Prairie Chicken 
IN Autumn 

Weights of 192 hunter-killed greater prairie chicken were ob- 
tained on October 24, 1951, by the use of spring balances. The 
balances were calibrated in 25 gram intervals. The weights were 
analyzed and presented in the same fashion as were the data on 
lengths of primaries. See table 3 and figure 12 on weight. 


Table 3. Weight Characteristics of the Greater Prairie Chicken (cf. 

Figure 12). Headings as in Table 2. 

in X±2— Sx V n 

Sample Weights " 

Males 700-1100 951.20 ± 10.48 52.92 5.56 101 

Females 650- 950 807.30 ± 11.38 54.25 6.72 91 

Adults 750-1100 911.30 ± 16.12 81.70 9.00 103 

Young 650-1050 853.00 ± 19.88 93.75 10.99 89 

Old males 775-1100 975.40 ±: 10.62 39.75 4.07 56 

Young males 700-1025 926.40 ± 18.32 61.50 6.63 45 

Old females 750- 950 834.90 ± 11.82 40.50 4.85 47 

Young females 650- 875 777.90 ± 15.68 52.00 6.68 44 

The average weight of the males, 951 grams, is significantly 
greater than that of the females, 807 grams, and the average weight 
of the adults, 911 grams, is significantly greater than that of the 
young, 853 grams. Furthermore, each sex- and age-class (old fe- 









I 1 1 ' 1 ' ' ' 1 

600 700 800 900 1000 MOO 


Fig. 12. Variation in the weight of the greater prairie chicken by 
sex and age (cf. Table 3). Diagrams prepared as for Figure 23. 


males, old males, young females, and young males) has an aver- 
age weight that is significantly different from that of each of the 
others, but there is a broad overlap between classes. The difference 
between the weights of birds of the year and birds more than one 
year old indicates that the young have not attained full size in late 
October. The low variability of both weights and lengths of pri- 
maries in young birds strengthens the conclusion that most of the 
young are hatched within a short period of time. 



700 800 900 1000 1100 gm». 



Fig. 13. Weight of adult greater prairie chicken (X axis) plotted 
against the length of primary number eight (Y axis) and the 
regression line of Y on X. 

Figure 13 illustrates the correlation between the lengths of pri- 
maries and the weights of individuals in adult birds. This figure 
was prepared from the same data from adult birds that were used 
in the preparation of Tables 2 and 3 and Figures 11 and 12. 
Adult males could be distinguished from adult females in most 
instances if both the weights and lengths of primaries were known. 
The impossibility of obtaining such data from hunters on a mass 
basis makes the determination of sex impractical by this means. 

Food Habits of the Greater Prairie Chicken 

Studies of foods eaten by the greater prairie chicken show that 
cultivated grains make up an important part of its diet. Judd 
(1905) examined 71 stomachs from the Midwest and Canada, rep- 
resenting each month except July, and reported the following foods: 
insects 14.11 per cent, grain 31.06 per cent, flowers and shoots 


25.09 per cent and fruits 11.79 per cent. The fruits were mostly 
rose hips, but included hazelnuts and the acorns of scarlet oak. 
Hamerstrom, Hopkins and Rinzel (1941:185) list cultivated grains, 
weed seeds and the browse of trees as the important foods in winter 
in Wisconsin. Schwartz (1945:72) found little evidence of the use 
of browse of trees in Missouri, but com and sorghum were im- 
portant winter foods. Yeatter (1943:414) reported ninety-one per 

cent plant food and nine per cent insects in summer in Illinois. 
No reports, relating restrictedly to the Midwest, have been found 

listing foods used by prairie chickens prior to the growing of cul- 
tivated crops there, but the early writings of Audubon (1834:491- 
501) and Koch (1836:159) in reference to the prairie chicken in 
Kentucky and Missouri, respectively, list fruit, tree buds, grain, 
wild grapes, mistletoe, buds of willow and hazelnut, seeds of grass 
and flowers (presumably forbs generally), and the catkins and 
shoots of hazelnut. The German naturalist, Koch (loc. cit.), de- 
scribed conditions in the state of Missouri soon after it was settled, 
and mentions that the "cupido-huhn" moved from the prairies to 
the cultivated land along the Missouri River in autumn. 

In the present study, information concerning foods and feeding 
habits of the greater prairie chicken was obtained by observing the 
birds in their feeding activities, by examining crop- and gizzard- 
contents and by studying droppings found in the Welda Area. 

Crop- and gizzard-contents were washed, dried and analyzed 
by separating the food items and measuring the volume of each 
kind of food. Samples made up mostly of insects were preserved 
in alcohol and measured in a moist condition. Droppings were 
collected as entire deposits from a roost, and as such would contain 
remains of most, or all, of the foods eaten late in the previous day. 
Only fresh droppings were collected in order to make certain of the 
date of deposition. Droppings were washed free of urates and 
foreign material and then dried. The remains of the various food 
items were identified and an estimate was made of the percentage 
of each. The total volume of each sample was measured and the 
volume of each item was computed from the estimated percentage. 
This procedure is less exact than the analysis of crop- and gizzard- 
contents, but does yield a broad picture of the trends of feeding 
habits throughout the year that cannot be obtained otherwise, with- 
out sacrificing many birds. Sixty-five samples of droppings, 29 
crops and 20 gizzards were analyzed. 

In the droppings the remains of the more common seeds could 
be classified and insect material usually could be classified to 
family. Dicotyledonous leaves usually retained their shape, and 


the various cultivated legumes were readily identified. Leaves of 
grass seemingly were more thoroughly digested, but could be 
identified as such when found in droppings. 

The greater prairie chicken ordinarily feeds early in the morning 
and late in the afternoon. In winter the usual behavior of the flock 
intensively studied near Welda was to fly to and from the feeding 
area, but some flocks were observed to walk to adjoining loafing 
cover from the feeding area. A female that was feeding on the 
leaves of weeds pulled off whole leaves approximately three inches 
long. Captive birds fed in similar fashion on lettuce leaves and 
swallowed in one piece leaves that were estimated to be 10 square 
inches in area. Whole seed-heads of crested plantain, Plantago 
aristata Michx., and parts of seed heads of sorghum with several 
seeds attached were found in crops. The manner of "browsing" is 
much like that of the turkey; the food is grasped and removed from 
the plant by an upward pull. Greater prairie chickens were never 
observed to scratch or otherwise attempt to uncover food on the 
ground, nor were they observed to browse in trees, although they 
were seen in trees in the winter of 1949-1950. 

Habit, or an attachment to a known feeding area, seems to govern 
the selection of a feeding place. In the winter of 1950-1951, F-2 was 
used by 50-60 prairie chickens and a number of hogs and cattle. 
By spring, feed seemed to be gone, yet the birds continued feeding 
there while the same foods were abundant at F-1. The birds were 
observed to thresh soybeans from the pods at this time of scarcity. 
In the autumn, entire seed pods of soybeans were found in crops. 

The data obtained from the analysis of crop- and gizzard-contents, 
from birds taken in the hunting season, are summarized in Table 4. 
The bulk (74 per cent) of the food was seeds and leaves of cul- 
tivated crops. Only 18 per cent was other plant material. Weed 
seeds (8 per cent) probably were used as much in October as in 
any other month. 

Of the greater prairie chickens collected in July and August, 
1951, three adults and five juveniles had food items in the crop or 
gizzard. Only a trace of insect remains was found in the adults, 
but in juveniles, insects made up approximately 25 per cent of the 
total volume of food, and occurred in each crop and gizzard that 
contained food. One individual had fed almost entirely on the 
larvae of a noctuid moth. The most important food item in summer, 
found in crops and gizzards of both adults and young, was oats; 
second in importance were the leaves of Korean lespedeza and 
other dicotyledonous plants. Seeds of four species of weeds were 


Table 4. Food Items Found ent 29 Crops and 20 Gizzards of the Greater 
Prairie Chicken Taken in October 1950 and 1951, from Anderson, Wood- 
son AND Coffey counties, Kansas. 

Per cent 

Food Item 







Panicum sp 










Tritictim aestivum ... 




Sorgum vulgare 


en 1 o 





J /\ /% 



111 pnf^ 1 pH nn pji p 

T Tniripnl'iripri 








Rosa sp 

. . s 




Soya max 




Lespedeza stipulacea 




Lespedeza stipulacea 


.... L 



Strophostyles leiosperma 










^ p.dir.fipn xfitii'tn. 









n n't 1 Q ci*\ 




Vitis sp 



A 1 A 



Abutilon theophraste 




Hibiscus trionum 



Cornus paniculata 












SumuhoricariDus orhiculatus 




Ambrosia trifida 




A. bidentata 




Ambrosia sp 




Helianthus annuus 




Plant debris 





Locustidae adults and 





Table 4. Concluded 

Food Item 

Per cent 

Consumed Occmrence Volume 


Coccinellidae adults 


Pentatomidae adults 


Cicadellidae adults 

Lepidoptera larvae 










Total plant material ( per cent of total volume ) 

Cultivated crop seeds 

Cultivated crop leaves 

Wild forb leaves 

Grass leaves 

Weed seeds 


Total insect material 
Total grit 



In the column headed "Part consumed," the letter "S" represents seeds and 
"L" represents leaves. Hard seeds such as those of Rosa and Cornus were found 
only in the gizzards and wear on seeds suggested that they functioned as grit. 







I I 


Fig. 14. Seasonal foods of the greater prairie chicken in Kansas. 
Cultivated grains are staple foods in winter and green foods are 
used extensively in winter. 

found, but those of no one made up so much as one percent of the 
total volume of food and seeds of weeds obviously make up only a 
small part of the food under modem conditions. 


All food items found to have been used by the greater prairie 
chicken are listed in Table 5. To ascertain the foods eaten, study 
was made of one or more crops or gizzards in each of the months 
of July through December and droppings were studied in each of the 
twelve months. The combined data on food are shown in Figure 14. 

Table 5. Monthly Occurrence of Food Items Found in Droppings, 

Crops and Gizzards. 

Food item consumed JFMAMJJASOND 


Gramineae leaves xxxxxxxxxxxx 

Panicum sp seeds x x x 

P. dichotomiflorum . . . , seeds x 

Avena sativa seeds x x x x 

Zea mays seeds xxxxx xxxx 

Sorgum. vulgare seeds xxxx xxx 

Triticum aestivum seeds x x 


Cyperus sp seeds x 

Scleria triglomerata .... seeds x 


(other than cultivated 

legumes) leaves xxxxxxxxxxxx 


Polygonum sp seeds x x 


Rosa sp seeds x x 


Soya max seeds x xxx xxx 

Lespedeza stipulacea . . . seeds x x 

Lespedeza stipulacea. . . leaves x xxxxxxx 

Trifolium repens leaves x x x 

Medicago sativa leaves x x 


Oxalis sp seeds, pods x 


Rhus sp seeds x 


Vitis sp seeds x 


Hibiscus trionum seeds xxxxx 

Abutilon theophraste. . . seeds x 


Cornus sp fruits x x 


Table 5. Concluded. 

Food item cons^umed JFMAMJJASOXD 


Ruellia sp seeds, pods, 

calices. ... x 


Plantago arisiata seed heads x 


Diodea teres seeds x x x 


orbiculatus seeds, berries x x 


Ambrosia trifida seeds x 

A. hidentata seeds x x x x x x x 

Ambrosia sp seeds x x 

Helianthus annuus .... seeds xxx xxxx 


Phasmidae entire x x 

Locustidae entire x x 

Coleoptera (unidentified) entire x x x x x x 

Chrysomelidae entire x x 

Coccinellidae entire x 

Carabidae entire x 


(Pentatomidae) entire x x 


In autumn, winter and spring, the greater prairie chicken fre- 
quently feeds on wheat. Much of the material found and identi- 
fied as leaves of grass probably was wheat. In eastern Kansas 60 
to 70 per cent of the food consumed by the greater prairie chicken 
is thought to be derived from cultivated crops, either seeds or 
leaves, whereas insects and weed seeds make up approximately 5 
per cent and leaves of forbs and wild grasses make up the remainder. 

At first inspection, these data might seem sufiicient cause for 
condemning the greater prairie chicken as detrimental to agricul- 
ture. There is no doubt that the bird does consume large quan- 
tities of grain. Hamerstrom et al. (1941:193) found that penned 
birds of this species consumed, in addition to browse, approxi- 
mately one and one-half ounces of grain per day per bird. This 
seems to be a fair estimate for wild birds, for a full crop contains 
more than one-half ounce of grain, and a bird would presumably 
eat more than two full crops of food per day. Accepting this figure 
as the average amount of grain consumed per day, a flock of 100 
birds would consume approximately four and one-half bushels of 
grain each 30 days. In areas with flocks of 100 or more birds per 
square mile, as found in this study, it is obvious that a considerable 
quantity of grain would be consumed by them. 

But, what is the source of this grain? Modem agriculture is based 
on the use of machinery. It is economical for the farmer to use 
machinery to harvest his crops, even though other methods might 
result in less waste. Formerly, when most grain crops were shocked 
and left in the field until threshed or fed, prairie chickens and other 
species made inroads upon this grain. In the Welda Area most 
crops were combined from the standing grain, or the shocks were 
removed from the fields before prairie chickens had begun to 
feed from them. An exception is that some oats remained in the 
shock during the wet weather of 1951. Mechanical harvest of crops 
in the field, leaves feed unavailable to cattle. This waste feed was 
the source of most of the grain used by prairie chickens in the 
Welda Area. At no time were the birds observed to use any 
grain other than waste. It is known, however, that damage to 
shocked grains occurs when the shocks are left in the field during the 
autumn and winter. This practice leaves the crop subject to loss 
from many granivorous forms of wildlife, and is not recommended. 

The second large item of cultivated crops used by the greater 
prairie chicken is the leaves of legumes. The most commonly used 
cultivated legumes are those found in pastures — Korean lespedeza 
and white clover. Intensive feeding on newly sown alfalfa would 


probably be detrimental, but alfalfa was the least used of the 
cultivated legumes. Large numbers of greater prairie chickens do 
not occur in Kansas where less than two-thirds of the land is in 
native grass. Under such conditions, prairie chickens could hardly 
cause an appreciable loss of forage that was intended for cattle. 

Fortunately, these facts are recognized by many farmers in areas 
where the greater prairie chicken occurs. These farmers recognize 
that much of the grain eaten by prairie chickens would otherwise 
be wasted. Even in 1949, when the population of the greater prairie 
chicken was high, few farmers interviewed complained of damage 
by it. Most farmers were interested in the welfare of the greater 
prairie chicken or at least tolerant of it. 

There has been a tendency on the part of some authors to justify 
the presence of birds on the basis of the number of insects, detri- 
mental to agriculture, consumed. This justification has had much 
to do with the widespread acceptance of birds as desirable. It is 
true that, in the aggregate, birds consume large quantities of insects. 
It is equally true that insects are adapted, by their reproductive 
capacity, to furnish this food for birds and other predators, with- 
out their numbers being unduly reduced. Lack (1951:443) points 
out that birds should not be regarded as harmful or beneficial unless 
the eflFects of birds on their food supply are known. In the Hght of 
present knowledge, it seems that we should regard any one species 
as a part of the integrated web of living things (the biota), and 
evaluate each species according to its net worth, whether this value 
be economic, esthetic or recreational. Viewed in this perspective, 
prairie chickens are eminently desirable, for they possess great 
esthetic appeal as part of a greatly diminished prairie biota, they 
provide a source of recreation for many hunters and are at least 
neutral in relation to agriculture as now practiced. 

Population Changes of the Greater Prairie Chicken 

A preliminary report (Baker, 1952) summarizes a part of the 
information gathered in this study concerning population changes 
of the greater prairie chicken in Kansas. According to Stene 
(1946:9), prairie chickens were given protection by the first Kan- 
sas Legislature in 1861. This act, providing a closed season on 
prairie chickens, partridge and turkey, presumably reflected a deple- 
tion that had already occurred in the numbers of these birds. Com- 
mon carriers were made liable for transporting illegal game in 1876. 

The year 1903 marks the first time that an open season was not 
held throughout the State. The major points in the regulation of 


the hunting of prairie chickens in Kansas since 1903 are summarized 
as follows; 

1903 — season closed in some western counties and two eastern counties for 
the first time, 

1913 — season closed throughout the State for the first time, 

1921 — season reopened September 20 to September 30, daily limit of five, 

1925 — season closed, 

1931 — season reopened for two days, 

1935 — season closed, 

1941 — season reopened, one day, in six eastern counties, 

1942 — season opened, one day, in eight eastern counties, 

1943 — season opened, two days, in eight eastern counties, daily limit three, 
possession six, 

1944 — season closed, 

1950 — season reopened, one day, in fifteen eastern counties, limit two, 

1951 — season opened one day, in sixteen eastern coimties, limit two, 

1952 — season opened one day, in eighteen eastern counties, limit two. 

In 1943, few hunters succeeded in taking their hmit, and many 
were completely unsuccessful. It may be inferred from this that 
a marked decrease in numbers of prairie chickens occurred in the 
period 1941-1943. At the beginning of this investigation, in 1949, 
the greater prairie chicken was again abundant, indicating a marked 
rise in numbers from 1943 to 1949. A conservative estimate is that 
there were 50,000 greater prairie chickens in Kansas in 1949. 

Information concerning population changes of the greater prairie 
chicken was obtained in this study by repeated censuses of the 
Welda Area, particularly of one flock, and by interviewing hunters 
in the open seasons. Repeated counts of males using flock range A 
(see Figure 5) were made each fall and spring, and less frequent 
counts of the birds using flock range C were made. In each open 
season, birds were examined in the field to learn the sex and age 
of greater prairie chickens bagged, and hunters were interviewed 
to learn the number of birds killed and the length of time spent 
in hunting. In 1950, four investigators accompanied state game 
protectors and examined greater prairie chickens killed in six coun- 
ties. In addition the writer concentrated his efforts in this respect 
in the Welda Area. In 1951, four two-man teams worked inde- 
pendently of the state game protectors in four counties characteristic 
of different types of habitat of the greater prairie chicken. In 
1952, three two-man teams and two individuals working alone 
gathered similar data in six counties. 

All greater prairie chickens examined were classified as young of 
the year, or as adults, by noting the condition of the two outer 
primaries ( see Ammann, 1944 ) . Sex was ascertained by examining 


the coloration of the rectrices, the pinnae and the air sacs ( Ridgway 
and Friedmann, 1946:206). 

The 1949-1952 Decline in Numbers 

The changes in the population on the study area are presented 
in Table 6. These data represent the highest number of birds 
present in each flock for any one season in one day. No data were 
obtained from flock range C in the autumns of 1949, 1950 or 1951. 

Table 6. Number of Greater Prairie CracKENS Using Two Flock 
Ranges on the Welda Area, 1949-1952.. 

Season and Year 

Flock Range 

A C 

Fall of 1949 146 ? 

Spring of 1950 104 43 

Fall 'of 1950, before hunting 145 ? 

Fall of 1950, after hunting 72 ? 

Spring of 1951 47 31 

Fairof 1951, before hunting 42 ? 

Fall of 1951, after hunting 24 ? 

Spring of 1952 15 15 

Fall>f 1952, before hunting 8 18 

Fall of 1952, after hunting 6 12 

Data on sex and age of greater prairie chickens examined in the 
two open seasons are presented in Table 7. Anderson and Woodson 
counties are representative of the best range of the greater prairie 
chicken in the Eastern Area. Chase, Chautauqua, Butler and 
Cowley counties are typical of the Bluestem Hills. The sample 
from Wabaunsee County was taken at the edge of the northern part 
of the Bluestem Hills and is representative of a condition inter- 
mediate between the Bluestem Hills and the Eastern Area. Data 
were obtained from 273 birds in 1950, 212 birds in 1951 and 119 
birds in 1952. 

Fluctuations of the population of greater prairie chickens in 
Kansas, prior to 1913, are not demonstrable, unless the action of 
the State Legislature in 1903 in closing some counties to hunting 
be taken to indicate a low point in numbers at that time. An all- 
time low seems to have been reached in 1913, when the season 
was closed throughout the State for the first time. A period of 
abundance in 1921 is reported by Clapp (1922:9). If open seasons 


indicate high populations, peaks of abundance occiured at the 
turn of each decade since 1920. Table 6 indicates that there was a 
marked reduction in the number of greater prairie chickens on the 
study area in the course of this study. 

The reduction in numbers probably was not so great throughout 
the entire area open to hunting as it was in the Welda Area. It was 
estimated that in 1950 there were 30 to 35 hunters per square mile 
in the vicinity of Welda, and almost as many in 1951, but fewer in 
1952. However, in adjacent Linn and CojBFey counties in 1950, 
game protectors reported few hunters. Even in northern Anderson 
County, a more moderate decline in numbers is revealed by counts 

Table 7. Frequency of Occurrence of Sex- and Age-Classes of Greater 
Prairie Chickens Examined in 1950, 1951 and 1952. 

Number of Prairie Chickens Examined 

County 1950 1951 1952 

Males Females Males Females Males Females 








































Chautauqua . . 












Greenwood . . . 





































of birds on booming grounds made in the spring of 1950 and in the 
spring of 1952. In 1950, there were 67 males on three booming 
grounds, and in 1952, there were 24 males on the same booming 
grounds. Even within and immediately adjacent to the Welda Area 
the effects of reduced hunting pressure were noted. The half- 
section pasture that was the principal range of flock C was closed 
to hunting in 1951. This flock suffered less loss than did the birds 
of flock A. Furthermore, in the quarter section immediately east 
of the area of study, hunting was prohibited in 1951, and the birds 
using this quarter section declined from approximately 75 in the 


spring of 1950 to 32 in the spring of 1952, as compared to the change 
from 104 to 15 in flock A in the same period of time. 

Figure 15 shows the trends of the population of the greater 
prairie chicken in Missouri from 1920 through 1944, as given by 
Schwartz (1945:36), with the open seasons in Kansas superimposed 
by crosshatching. Also included is the hypothetical rise in the 
population of the greater prairie chickens in Kansas from 1943 to 
1949 and the decline from 1949 through 1952 as indicated in the 
above discussion. Residents of the Welda Area say that the greater 
prairie chicken was nearly as numerous in 1948 as in 1949. Figure 
15 reveals that the open seasons in Kansas have coincided roughly 
with the high populations of Missouri, but that at least the open 
season of 1941 fell somewhat after the peak population was attained 
in Missouri. Furthermore, the three consecutive open seasons, 1941- 
1943, were accompanied by a more pronounced decline than oc- 
curred in Missouri, without hunting, in the same period. 

1920 1925 1930 1935 1940 1945 1950 





Fig. 15. Population trends of the greater prairie chicken in Mis- 
souri (after Schwartz, 1945). Open seasons in Kansas super- 
imposed by crosshatching, the population curve from 1941 to 1949, 
inclusive, is estimated for the state of Kansas, and from 1949 to 
1952, inclusive, is from the original data for the state of Kansas. 

It should be emphasized that heavy natural losses of adult greater 
prairie chickens occurred in the period 1949-1952 in addition to 
the loss from hunting. Were this not true, the kill of approximately 
50 per cent of the population each year, as indicated by the numbers 
recorded before and after hunting in Table d, would have been 
balanced by the productivity of approximately 50 per cent, shown 
in Table 7, and the population would have remained more nearly 
static. It is evident, therefore, that the 1950, 1951 and 1952 hunt- 
ing occurred in a period of naturally dechning population, or at 
best a stable population and that the 1941-1943 open seasons prob- 
ably occurred under similar conditions. It might be postulated 
that if open seasons were held in a period of increasing population, 


when production of young exceeds natural mortality, this popula- 
tion could sustain a greater kill by hunters without undue deci- 
mation. If this be true, a more efficient utiHzation of the prairie 
chicken as a resource could result, and high populations, which 
may elicit complaints from farmers and ranchers, could be avoided. 

Changes in Age Composition 

The age ratios, as revealed by the examination of birds killed 
by hunters (see Table 8), reflect the weather conditions of the 
three breeding seasons, and the average size of broods seen in 1950 
and 1951 (cf. Figures 7 and 8). The productivity of the greater 
prairie chicken, as shown in Table 8, falls far short of that of the 
bobwhite (Bennitt, 1951:32), which experience has shown is a 

Table 8. The 0.9546 Confidence Limits of Certain Population Char- 
acteristics OF THE Greater Prairie Chicken, From Birds Examined in 
THE Autumn, 1950, 1951 and 1952. 

Sex and Age 


Mean and Limits 


Per cent of females 



0.414 ±0.089 

Per cent of young 




Per cent of females 

among young birds . . 



0.393 ±0.125 

Per cent of females 

among old birds 




notably successful game bird. Yet the productivity of the greater 
prairie chicken found in my study is similar to that found by Lee 
(1950:476) for the lesser prairie chicken in New Mexico in 1949. 
If further study should reveal that 50 per cent is near normal pro- 
ductivity for the greater prairie chicken, this fact should be con- 
sidered in its management. Under the conditions now existing in 
Kansas, no species with such a low productivity can withstand 
heavy hunting eaclf year. The history of prairie chickens in Kansas 
indicates that their productivity, in view of their present limited geo- 
graphic range, is not sufficient to provide annual moderately re- 
stricted hunting, for in each of the decades since 1920 open seasons 
have been held, and in each instance thereafter the population de- 
chned to so such a low point as to necessitate the closing of the 
hunting season for a number of years. 


Changes in Sex Composition 

Few data have been published relative to the sex ratios of prairie 
chickens. Schwartz (1945:14) calculated sex ratios from birds seen 
on booming grounds and found that only approximately 32 per 
cent of the population were females. Davison (1940:58) found 
a similar sex ratio in young of the lesser prairie chicken, and Lee 
(1950:477) found, in the same species, a preponderance of young 
males and old females in the bags of hunters in New Mexico. 

The percentages of females, of the greater prairie chicken, killed 
by hunters in 1950, 1951 and 1952 in Kansas are given in Table 8. 
Of nineteen captive chicks, from the Welda Area, nine were of 
one sex and ten of the other, indicating a sex ratio at hatching of 
near 50:50. Significant departures from this ratio occurred among 
old females in 1950, but among young females the departure was 
not significant for the size of the sample, but it was suggestive 
of a differential mortality among young in favor of the males. The 
sex ratio did not depart significantly from 50:50 in the sample ob- 
tained in 1951 or 1952. It is not possible to say whether these dif- 
ferences are real or result by chance from sampling, but there 
seems to be but little doubt that, on the average, there are more 
males than females of the greater prairie chicken. 

Differences in Abundance 
as Indicated by Hunters' Success 

The success of hunters as an index to relative populations of game 
has been used in Missouri since 1947 (see Crawford, 1951:307). 
The method is theoretically sound, because the number of game 
animals bagged in a given length of time, on the average, should 
be related to the density of the population of game. 

In each hunting season ( 1950, 1951 and 1952 ) , hunters were in- 
terviewed to ascertain the number of birds killed and the time 
spent in hunting. The resulting data were computed in terms of 
birds killed per gun hour, and are presented in Table 9. The dif- 
ferences between each area sampled and between each year are 
highly significant statistically in most instances. To test the value 
of data on success of hunters we may first compare the results of 
the three hunting seasons in the Welda Area where populations 
were known. In 1950, hunters in the Welda Area killed, on the 
average, 0.376 birds per gun hour, and in 1951, 0.128 birds per 
gun hour. In 1952, too few birds were killed in the Welda Area 
to permit comparison. These data compare favorably with the 
populations of 145, 42 and 8 for 1950, 1951 and 1952 respectively 


(Table 6). In other parts of Anderson County, populations prob- 
ably did not decline so much in 1951 and 1952, and the hunting 
success was 0.214 birds per gun hour, 0.124 birds per gun hour and 
0.059 birds per gun hour for 1950, 1951 and 1952 respectively. 

A second test of the value of the success of hunters as a measure 
of relative populations may be made by comparing the data from 
Anderson and Woodson counties. The relatively poorer success 
in Woodson County, in 1951, may be correlated with the poorer 
productivity in that county, in 1951, as indicated by the low number 
of young birds bagged by hunters ( see table 7 ) . There were only 
32.65 per cent of young in the sample from Woodson County, 
compared to 46.74 per cent in the sample from Anderson County. 
It is concluded that the average time required to bag a prairie 
chicken is a useful measure of relative densities of populations. 
On this basis, in the counties sampled, the greater prairie chicken 

Table 9. The Mean Number of Birds Per Gun Hour and the 0.9546 
Confidence Limits of the Means, 1950, 1951 and 1952. 

Mean and Limits 

County 1950 1951 1952 

Anderson 0.214=^0.001 0.124=^0.002 0.059^0.006 

Wabaunsee . 144 =fc . 016 . 134 =t . 002 . 271 ± . 024 

Woodson 0.199=^0.008 0.092±0.008 0.113±0.014 

Chase 0.082^ 0.005 

Cowley, Chautauqua and 

Butler 0.164 ±0.019 

State wide average . 185 . 109 . 109 

is least numerous in Chase County, a part of the Bluestem Hills, 
and there were fewer birds in 1951 than in 1950 throughout the 
entire range in Kansas. Furthermore, the population in 1952, as 
indicated by hunters' success, made Httle recovery in spite of dry 
weather in the hatching season. The state-wide hunting success 
was the same for the two years (see Table 9). It is especially im- 
portant to note that in areas of heavy hunting pressure, as in Ander- 
son County, hunting success declined in 1952, whereas in less popu- 
lar hunting areas (for example, Wabaunsee County), the success 
remained relatively good. 



In general, census techniques have to be adapted to the peculiari- 
ties of the species censused. Most methods previously proposed 
for the census of prairie chickens have been based on their mating 
behavior. A census, adapted to conditions in Kansas, of prairie 
chickens on their booming grounds is recommended. 

Two conditions found in Kansas should be recognized in out- 
lining a census based on booming-ground counts. First, high winds 
frequently interfere seriously with the observer in the locating of 
booming grounds by the sense of sound. Experience has shown 
that this diflficulty can be overcome, first, by hmiting the census 
strip to one-half mile either side of the line of travel, second, by 
frequent stops and the use of binoculars, and third by being alert 
for birds to be seen against the horizon. Second, many continuous 
square miles of the Bluestem Hills and Blackjack Prairies are 
without county roads; in such roadless territory a system of census- 
plots should be used in place of census-strips. The procedure ex- 
plained immediately below is modified from Schwartz (1945:31), 
and is recommended for censuses in spring. 

1. Select adequate census areas to sample the entire range of 
both species of prairie chickens. In areas where roads form a 
regular grid pattern, census-strips one-half mile either side of a 
road should be selected. In roadless areas, census-plots of at least 
four square miles should be used as recommended by Davison 
( 1940 ) . Experience indicates that six miles of strip or four square 
miles of plot should be covered in one morning. 

2. Preferably one census taker should be available for each 
county and should be assigned three or four census areas. In coun- 


ties where prairie chickens are hmited to only part of the county, 
proportionately fewer census areas could be assigned per county, 
and one observer might cover more than one county. 

3. On one or more centrally located booming groimds the per- 
son in charge of the census should observe the progress of events 
until it is evident that the peak of mating activity is occurring. This 
is indicated by the presence of a fairly constant and high number of 
females appearing on the booming ground and usually occurs 
early in April. The census should be conducted in the week fol- 
lowing the peak of mating. In the week of the census, the same 
booming ground should be observed to ascertain the number of 
females that are present on it each morning. 

4. The actual census should be conducted for two hours in the 
morning beginning as soon as there is suflBcient light to permit accu- 
rate observation. When possible, the counts should be made without 
flushing the birds and separate counts of males and females should 
be made. With a little experience, the census-taker readily can 
distinguish males from females by the more slender appearance 
and the attitude of indifference in the latter (Schwartz, 1944). 
Males ordinarily would be engaged in display, combat and pur- 
suit activities, or might be squatting quietly at times when no fe- 
males were present. With moderate to high winds, stops should 
be made each half mile on a roadside census, the surrounding 
terrain should be studied with the aid of binoculars, and any areas 
not visible from the road should be visited. When conditions for 
hearing are good, stops need be made at distances of only a mile, 
and visual survey of unseen parts of the census area is unnecessary. 
Complete coverage of each area is essential, or if an abserver can- 
not cover the assigned areas, the part of each that was censused 
should be designated. Preliminary surveys of the census areas 
to locate booming grounds would facilitate the census, provided 
that the census was made only in the assigned week, and provided 
also that sole reliance was not placed on the booming grounds that 
were found in the preliminary survey. 

5. All findings should be reported on standard forms to the 
person in charge of the census. The number of males per square 
mile would then be calculated. Total figures for the population of 
the entire state would not be necessary provided no major changes 
in the area occupied by prairie chickens occur. 

Considering the potential losses, among young birds, caused by 
adverse weather, spring censuses are not sufficient bases for fixing 
regulations. Such counts would be most useful in ascertaining the 


trends of the population and the breeding stock present at the be- 
ginning of the breeding season. The 1951 and 1952 data relative 
to the spring and fall populations demonstrate that there was no 
net gain in numbers through the period of reproduction. It is im- 
perative, therefore, to evaluate the success of reproduction each 
year in order to predict the fall population. A careful analysis of 
the amount and distribution of rainfall in the hatching period, May 
15 to June 15, supplemented by counts of the number of birds seen 
in broods from July 1 to August 15, should be sufficient to supple- 
ment the spring census in predicting the fall population. If such 
a procedure were used over a period of years, a correlation between 
weather conditions and reproductive success probably could be 
established. If so, the counts of broods might no longer be neces- 
sary. Counts of broods might be obtained by field men incidental 
to their other duties; but early morning censuses with the aid of 
dogs probably would be necessary to obtain sufficient data. An 
accurate census of the fall population on the booming grounds 
could be made as early as the first or second week of October. 

Hunting Regulations and Refuges 

With adequate censuses it should be possible to set open seasons 
to harvest the surplus prairie chickens most efficiently. If seasons 
were opened when the census indicates that the population of 
prairie chickens is increasing and approaching peak numbers, full 
advantage could be taken of the favorable reproduction, and exces- 
sive abundance of prairie chickens could be avoided. 

Judging by the efiFects of hunting in 1950, 1951 and 1952, more 
liberal regulations than those then appHed cannot be permitted un- 
less a much larger area is available for hunting, or unless there are 
fewer hunters. In the three seasons, in areas of high prairie chicken 
population, there were too many hunters. Too large a number of 
hunters placed excessive pressure on the prairie chickens, and also 
interfered with good sport. 

Regulating the number of hunters by issuing limited numbers of 
special permits would be possible, and might eventually become 
necessary. A permit system could be used to gain also a better 
distribution of hunters. Under present circumstances the permit 
system is thought to be unnecessary, provided other measures are 
taken to insure the survival of a sufficient breeding stock. 

One measure that should be seriously considered is to open 
the prairie chicken season at a time when the season is open on 
pheasants. This suggestion has been favorably received by land- 
owners both in the pheasant country and in the prairie chicken 


country, for both are aware of an undue concentration of hunters. 
There is no way of predicting the net result of such a measure, 
but it could be expected that hunting pressure would be reduced 
for both species at least when the seasons were concurrent. A 
disadvantage for the manager would be the lesser number of en- 
forcement personnel in any given area than there is now with the 
two seasons separated in time. 

SuflBcient protection probably could be given prairie chickens, 
with no limitation placed on the number of hunters, if a system of 
refuges were provided. It was pointed out in the section concern- 
ing population changes that small areas closed to hunting by land- 
owners were eflFective in reducing the kill in 1951. It was observed, 
especially in 1951, that more areas were closed to hunting by land- 
owners in areas where the numbers of the greater prairie chicken 
were small in Anderson County. In such areas, unsolicited posting 
of land probably will provide suflBcient refuge areas. In areas of 
high population of prairie chickens, where farmers may desire a 
reduction in numbers, few refuges of this nature are likely to be 
provided. Yet, such areas have the greatest concentration of 
hunters, and are in the greatest need of refuges. Active promotion 
of a private refuge system in areas with high populations of prairie 
chickens, with the aim of providing a minimum of four square 
miles of refuge per township, should provide suflBcient protection. 
Each refuge preferably should be one square mile or less in area. 

In places where fires in the spring commonly remove nesting 
cover over large areas, refuge or management areas are needed 
to provide suitable nesting cover. Much could be done in this 
respect by encouraging ranchers not to burn their grasslands, and 
by oflFering assistance in the protection of their lands against fire. 
Many small areas, perhaps a total of 40 acres per square mile, 
would function better than a few large areas, since prairie chickens 
do not range far. Serious consideration should be given to the 
acquisition and management of refuge areas in the Bluestem Hills 
and Blackjack Prairie, where extensive burning is most common, 
and where land values are relatively low. State-owned refuges 
could serve as nesting and winter feeding areas, as demonstrations 
of proper grazing practices, and could be at least partly self sus- 
taining by the lease of grazing rights. Special permits, similar to 
those used for quail hunting, could provide the funds necessary. 

Range and Pasture Management 

Prairie chickens are essentially birds of the grasslands. For- 
tunately, the practices recommended for the most profitable long- 


time use of grasslands are beneficial to prairie chickens. In much 
of the former range of the greater prairie chicken, however, where 
soils are adapted to the production of crops in rotation, the re- 
quirements of this species for permanent grass, probably can never 
be provided. In Kansas, the present acreage of native grassland 
is in Httle danger of reduction by plowing, for most of this land is 
unsuited to tillage. Every effort should be put forth, however, 
by agencies concerned with agriculture and conservation to pro- 
tect existing grasslands, and to encourage the re-estabhshment of 
permanent grass, especially native grass, on areas proven unsuitable 
for cultivation. 

Prairie chickens are benefited by moderate grazing of pastures. 
The paths and small areas of reduced cover resulting from the 
activities of cattle facilitate the movements of young birds, and 
provide places suitable for sunning in times when the grass is wet. 
Anderson (1946:95) recommends that grazing be deferred until 
four to six weeks after growth of grass begins in the spring, and 
that then the grass be grazed heavily for the remainder of the 
season. Under such a system, winter cover might be reduced by 
the heavy grazing, but spring burning would lose its appeal to the 
landowners, and nesting cover presumably would be more plentiful. 
Deferred grazing produces more pounds of beef per acre than early 
spring grazing and is well suited to the needs of the resident rancher 
who can provide supplementary pasture. 

The usual practice is to begin grazing approximately on May 
first. This is especially true of pastures leased for the grazing of 
cattle from the Southwest, in which case the desire of the cattlemen 
is to attain the most rapid and early gain possible. Cattle may be 
taken from the pastures as early as July first, and the growth of 
grass thereafter is not only wasted, but it interferes with the eflBcient 
utihzation of forage the next spring. The usual method of re- 
moving this unwanted growth is by burning. According to Ander- 
son (1946:100) there is some justification for burning since it in- 
creases the efficiency of grazing by eliminating the old growth and 
because it permits earlier grazing. Burning is not recommended 
by Anderson, however, because the earlier grazing is detrimental 
to the grass, and the total production of grass is reduced. So long 
as absentee ownership and the leasing of pastures are prevalent, 
there seems to be no cure for the harm done to upland game by 
burning, except through a program of education and a system of 
refuges. Ranchers who insist on burning should be encouraged 
to do so only when the ground moisture is plentiful and after a 
rain. Under such conditions some cover is left. 


Both deferred grazing without burning, and early spring grazing 
with burning have advantages for prairie chickens, but of the 
two, deferred grazing seems more desirable in that it provides * 
improved conditions for nesting by the greater prairie chicken. 


In Kansas, all known large areas suitable for prairie chickens 
have at least a few birds present. In general, animals are capable, 
by natural increase, of stocking new range as rapidly as it becomes 
suited to them. If, in the future, areas that seem suitable for prairie 
chickens are not naturally stocked, by reason of their isolation, 
stocking could best be done by trapping and transplanting wild 
birds. Care should be taken to trap sufficient females, because ex- 
perience has shown that they are more difficult to trap than are males. 

Summary and Recommendations 

1. Prairie chickens increased in numbers and extended their 
range with the development of early agriculture in the Midwest. 

2. With the intensffication of agriculture and with excessive 
hunting, the numbers and range of prairie chickens decreased. 

3. Since the time of the first reliable estimates of the numbers 
of the greater prairie chicken, this species has fluctuated regularly 
in numbers. In Kansas, the declines have been accentuated by 

open seasons. j 

4. Kansas is one of the four states having the largest number of 
greater prairie chickens. 

5. The greater prairie chicken in Kansas is confined to areas 
in which at least one-third of the land is in native grass, and is most 
abundant where approximately two-thirds of the land is in native 

6. The reproductive success of prairie chickens is low, because 
the reproductive period is short (late spring and early summer), 
and because adverse weather conditions may be detrimental to 
both eggs and young. The reproductive season seems to be the 
most critical period of the year for the species. 

7. Weights of the greater prairie chicken and lengths of its 
primaries are both different according to sex and within each sex 
according to age, but neither weights nor lengths of primaries pro- 
vide a sure means of distinguishing the sexes. Males average 
heavier than females. 

8. Young of the greater prairie chicken can be aged by the con- 
dition of the molt. 


9. In eastern Kansas, the greater prairie chicken feeds more on 
cultivated plants than on uncultivated plants. Of cultivated plant 
food eaten, most is waste that would not be harvested. 

10. The success of hunters provides a means of comparing 
density of population in different years and in different areas in the 
same year. 

11. A management plan is outlined for the greater prairie 
chicken, and is thought to be applicable to the lesser prairie chicken 
as well. The plan includes: 

A. Censusing the adult population in the spring and in the fall 
to ascertain trends in numbers; 

B. Censusing young birds between July 1 and August 15 to 
ascertain the success of reproduction; 

C. Providing open seasons only in periods of increasing num- 
bers, as indicated by the spring censuses, and in years of favorable 
reproduction, as indicated by the summer censuses; 

D. EstabHshing refuges 

(1) One square mile in each township throughout the Blackjack 
Prairies and Bluestem Hills. Preferably state-owned. 

(2) One section in each nine sections in the eastern parts of the range 
where hunting pressure is excessive. Could be privately owned. 

E. Encourage the preservation of existing prairies and the re- 
seeding to native grass of lands proven unsuitable for cultivation. 


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Transmitted January 26 y 1953.